Full text of "Mammals of the Soviet Union"

See other formats

The present book, of the series Mam-
mals of the Soviet Union, Sirenia and
Carnivora, is the fourth volume to be —
published in English. Previously Volume
I — Artiodactyla and Perissodactyla,
Volume II, part 2 — Carnivora (Hyenas
and Cats) and Volume II, part 3 —
Pinnipeds and Toothed Whales were
published in 1988, 1992, and 1996
respectively. Sea Cows, Wolves and
Bears constituted the second in the
series to appear.

The original Russian book is so
large (982 pp.) that its publication in the
English translation format would have
resulted in a book of about 1400 pages,

-which was considered too unwieldy.
Therefore, it was decided to publish
-Volume II, Part 1 — Sirenia and Car-
nivora, in two roughly equal parts. The
first (Part la) consists of the Steller’s sea
cow, the now-extinct marine mammal
once found only around the Commander
Islands in the western Bering Sea, and
the Russian species of the wolf and bear
families. The second half (Part 1b) will
comprise the various members of the
weasel family, plus the introduced
American raccoon. As indicated in the
foreword to the English edition of
Volume I, the senior author and editor
of the series, Dr. Vladimir Georgievich
Heptner, died in 1975, and did not
complete the projected series. However,
work continues, and Russian volumes on
baleen whales, lagomorphs and dipodid
rodents are now published.

A final point: since the appearance
of Volume II, part 2, the Soviet Union
has disappeared. However, the series was
written in the context of the former
“union of republics,” and it would be
confusing to re-edit the text to conform
to the current political reality.

Ta,
x Ne i
hapa

ЗЕ

Mammals of the Soviet Union

Volume II
Part la

MAMMALS OF THE SOVIET UNION

In Three Volumes

| Edited by
V.G. Heptner and N.P. Naumov

Vysshaya Shkola Publishers
Moscow, 1967

Mammals of the
Soviet Union

Volume II, Part la

SIRENIA AND CARNIVORA
(Sea Cows; Wolves and Bears)

V.G. Heptner, N.P. Naumov, P.B. Yurgenson
A.A. Sludskii, А.Е. Chirkova and A.G. Bannikov

Illustrators
A.N. Komarov and N.N. Kondakov

Scientific Editor
ROBERT S. HOFFMANN

Smithsonian Insitution Libraries
and
The National Science Foundation
Washington, D.C.
1998

Mlekopitayushchie Sovetskogo Soyuza
In Three Volumes
V.G. Heptner and М.Р. Naumov, editors

Vysshaya Shkola Publishers
Moscow, 1967

Translators: Bolos Abdul Malek Botros, Hamed Tantawi, Hosni Ibrahim
Youssef, Ali Abdul Moneim Moussa

General Editor: Nawal Mehallawi, Al-Ahram Center for Scientific
Translations, Cairo, Egypt

Library of Congress Cataloging-in-Publication Data
(Revised for volume 2, pt. Та)

Geptner, V.G. (Vladimir Georgievich), 1901—1975.
Mammals of the Soviet Union.

Translation of: Mlekopitaiuschie Sovetskogo Soiuza.
Bibliography: v. 2, pt. la, p.
Vol. 2, pt. 1b has index.
Supt. of Docs. no.: Si 1.2: Ar 7/2
Contents: v. 1. Artiodactyla and Perissodactyla—
v.2. —pt. la. Sirenia and Carnivora (Sea Cows ; Wolves and Bears)
у.2. —pt. 2. Carniyora (Hyaenas and cats).
1. Mammals — Soviet Union. I. Nasimovich, A.A.
II. Bannikov, Andrei Grigor’evich. Ш. Hoffmann, Robert S.
IV. Title.
QL728.S65G4713 1988 599.0947 85-600144

Published for the Smithsonian Institution Libraries
by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110 001

Printed in India at Baba Barkha Nath Printers, New Delhi

Foreword to the English Edition

The Smithsonian Institution Libraries, in cooperation with the
National Science Foundation, has sponsored the translation into
English of this and hundreds of other scientific and scholarly studies
since 1960. The program, funded with Special Foreign Currency under
the provisions of Public Law 480, represents an investment in the
dissemination of knowledge to which the Smithsonian Institution is
dedicated.

The present book of the series Mammals of the Soviet Union, Sirenia
and Carnivora, is the fourth volume to be published in English. The
project was initiated in 1975 as my contribution to joint U.S. —U.S.S.R.
studies on Holarctic mammals, sponsored by the Environmental Agree-
ment between the two countries. Previously, Volume I, Artiodactyla
and Perissodactyla, Volume II, part 2, Carnivora (Hyenas and Cats)
and Volume П, part 3, Pinnipeds and Toothed Whales were published
in 1988, 1992, and 1996 respectively.

Sea Cows, Wolves and Bears constituted the second in the series
to appear. The fact that this volume was not published in English
translation in the proper sequence is due to the fact that the translation
was performed under a different contract from that supervised by
Amerind Publishing Company, New Delhi, India. Moreover, the origi-
nal Russian book is so large (982 pp.) that its publication in the
English translation format would have resulted in a book of about
1400 pages, which we considered too unwieldy. Therefore, it was .
decided to publish Volume II, Part 1, Sirenia and Carnivora, in two
roughly equal parts. The first (Part la) consists of the Steller’s sea
cow, the now-extinct marine mammal once found only around the
Commander Islands in the western Bering Sea, and the Russian spe-
cies of the wolf and bear families. The second half (Part 1b) will
comprise the various members of the weasel family, plus the intro-
duced American raccoon.

As indicated in the foreword to the English edition of Volume I,
the senior author and editor of the series, Dr. Vladimir Georgievich

Heptner, died in 1975, and did not complete the projected series.
However, work continues, and Russian volumes on baleen whales,
lagomorphs and dipodid rodents are now published. It is hoped that
English translations of these later volumes will follow more promptly
than has been true of the first volumes of the series.

English readers interested in Dr. Heptner’s contributions as a mam-
malogist should refer to the foreword to the English edition of Volume
I. Conventions used in rendering geographic names, first stated there,
are reprinted here for the convenience of the reader. Geographic names
are generally transliterated directly, but a few exceptions were permit-
ted (e.g. Moscow instead of Moskva, translation rather than
transliteration of certain modifiers of place names, such as Northern,
rather than Severnaya, Dvina). Soviet administrative units are numer-
ous, and the following equivalents were employed in translation: krai,
territory; oblast’, district; raion, region; guberniya ( archaic), gover-
nance, uyezd, county. Also, in the original Russian text, rivers, mountain
ranges, and cities are often not explicitly identified, the Soviet reader
being presumed sufficiently familiar with the geography of the coun-
try to be able to understand from the context of the sentence what sort
of place is referred to. Complicating the matter is the lack of articles
as parts of speech in Russian. To assist the English reader, I have
adopted the following conventions: if a river is referred to, an article
precedes it; if a mountain range is referred to, it is translated as a
plural; if a city is referred to, it is singular, and lacks the article.
Examples are: the Ural (river); the Urals (mountains); Ural’sk (city).
Geographic place names are also inflected in Russian, and these have
been simplified by omitting transliteration of the inflected ending. For
example, the Russian phrase v Yaroslavskoi i Kostromskoi oblastyakh
is translated “in the Yaroslavl and Kostroma districts.” In cases where
the nominative form of the place name has an -sk ending this is,
however, transliterated (e.g., Omsk); when a Russian “soft $121” is
employed in a place name, this is transliterated as an apostrophe (e.g.
Khar’kov); the “hard sign” is rendered as a double apostrophe. Be-
cause of the large number of place names in this volume, it was not
possible to verify all of them, and some inconsistencies are likely to
occur. I would appreciate it if readers would bring any errors they
may notice to my attention.

In Volume II, part 1, I have also attempted to improve certain
aspects of the translation in order to reflect more faithfully the original
Russian text. For example, what was translated as “taxonomy” in

Vii

volume I is now in some places rendered as “systematic position;”
what was “economic importance” is now “practical significance,” etc.

One further point of confusion not apparent to me when Volume
I was translated also required clarification; that is the English trans-
literation of the senior author’s surname. This begins with the fourth
letter of the Cyrillic alphabet, which usually has a “G” sound in Rus-
sian. However, the surname was originally German (as indeed was
mine) and in the original German began with the letter “H” of the
Latin alphabet. Since Cyrillic has no equivalent of “H” this is usually
transliterated into “G” in Russian. However, I know from conversa-
tions with him that Dr. Heptner preferred to use the original Germanic
form of his surname rather than the transliterated version, which is
rendered as Geptner. The rules of transliteration employed by the
Library of Congress do not permit such flexibility, and the attentive
reader may notice that Library of Congress cataloging employs the
latter.

A final point: since the appearance of Volume II, part 2, the
Soviet Union has disappeared. However, the series was written in the
context of that former “union of republics,” and it would be confusing
to re-edit the text to conform to the current political reality.

Robert S. Hoffmann

National Museum of Natural History
Smithsonian Institution
Washington, D.C.

Foreword

5 The second volume of the “Mammals of the Soviet Union” is devoted
to descriptions of the terrestrial Carnivores (Order Carnivora), and the
sea cows (Order Sirenia), consisting in our fauna of only one recently
extinct species. There is a great deal of information in our country
about carnivorous mammals, which include the most important fur
animals. In the last 10 years they [furbearers] have been studied in
especially great detail. Various unpublished data of importance avail-
able to the authors were included in the descriptions. Therefore, the
size of this book is considerable.

All species in the indigenous families of the Order Carnivora are
described with the exception of the cats (Felidae) and hyaenas
(Hyaenidae), i.e. the suborder Feloidae. Descriptions of these excluded
families will follow in the next volume. In addition, the book includes
two acclimatized American species of carnivores—raccoon and mink.
The striped skunk, as is known, failed to be acclimatized in our
country and is therefore not included.

The classification of this order follows Simpson (1945). Simpson’s
orders themselves are, as is known, conventional. Only one exception
is admitted: pinnipeds, contrary to the opinion of Simpson and some
others, are considered as an independent order. The arrangement of
orders in the book follows the sequence from most specialized to the
most primitive, that is to say, from up to down. Within the orders in
this volume — as in volume I —the taxa and species are arranged,
according to their specialization. In the sequence of orders described,
however, one other deviation was made: carnivores (Carnivora), and
not pinnipeds (Pinnipedia) were placed after the sea cows, in order
that pinnipeds and whales could be included in one volume. For ori-
entation to the actual relationships of orders, a macrosystemic scheme
of the class is given [pg. xix].

All the characteristics of a group are presented in a standard for-
mat and deviation was permitted in only a few special cases. Group
characteristics, given in an extremely condensed form, are applicable

Хх

not only to the species of our fauna but to the entire group. Principal
attention is paid to the distribution and systematic composition of
groups, and the phylogenetic relationships of our forms within the
group. Paleontological information is very condensed, mainly accord-
ing to Simpson (1945), but partly from Thenius and Hofer (1960),
“Fundamentals of Paleontology” edited by Yu.A. Orlov (1962), and
various other sources.

In the species descriptions, attention was mainly concentrated on
geographic distribution and biology. Ranges are broadly presented,
based on available data, and only limits are described. Habitats within
these ranges are not mentioned, and only the most important marginal
localities are given. Historical changes in geographic ranges are traced,
and as much as possible the description is based on the “reconstructed”
range of the species, in depth for centuries of “historical” time. This
is, therefore, a suitable opportunity to establish a more or less actual
geographic area. A knowledge of this is obligatory for solving theo-
retical questions of zoogeography and for practical work with the
fauna’.

Ranges beyond the boundaries of our country—including those
“reconstructed’”—follow numerous works, but chiefly for critical veri-
fication, information provided by Seton-Thompson (1909—1910), Miller
(1912), Sowerby (1923), Anthony (1928), Aharoni (1930),
Nezabitowski (1934), Shortridge (1934), Phillips (1935), Heim-de-
Balsac (1936), Allen (1938—1940, 1939, 1942), Pocock (1939, 1941),
Cabrera and Yepes (1940), Hamilton (1943), Harper (1945), Simpson
(1945), Anderson (1946), Carter, Hill and Tate (1946), Prater (1947),
Tate (1947), Troughton (1948), Enar (1949)*, Calinescu (1951), Rob-
erts (1951), Ellerman and Morrison-Scott (1951), Morrison-Scott and
Hayman (1953), Bannikov (1954), Laurie and Hill (1954), Miller and
Kellog (1955), Siivonen (1956), Haltenorth and Trense (1956) Cabrera
(1958, 1960), Van den Brink (1958), Frechkop (1958), Markov (1959),
Misonne (1959), Hall and Kelson (1959), Hatt (1959), Dulic and Tortric
(1960), Burt and Grossenheider (1962), Burton (1962), Cockrum (1962)
and many others. Individual references to the above-mentioned au-
thors were not given in the text descriptions of ranges external to the
boundaries of the USSR. North American ranges were based in nearly

'More detailed information about reconstructed geographic ranges is available in
the work of V.G. Heptner. Dynamics of geographic ranges of some ungulates and
human factors. Sb. “Geographical Questions”, 1959. [Dinamika areala nekotorykh
kopytnykh 1 antropicheskii factor. Sb. “Voprosy Geographii,” 1959.]

* Not in Lit. Cit—Sci. Ed.

all cases on excellent maps in Hall and Kelson (1959), whose opin-
ions differed completely, in some cases, from the description of others.
All maps of geographic ranges are original, prepared by V.G. Heptner.

Unfortunately, we have not succeeded in achieving uniformity
among different authors in describing the biology of species. How-
ever, the general formai is identical to that encountered in the first
volume; only the description of the extinct species—the sea cow—
differs. Concerning the latter, instead of paraphrasing the data from
published literature, a special translation was made of the original
work of Steller — the only naturalist to have observed this animal and
studied its anatomy. Until now, i.e. for more than 200 years, Steller’s
text has not appeared in Russian. S. Krasheninnikov (1786), who had
Steller’s manuscripts, published a translation of only certain parts of
this work. Owing to the language in which the work of Steller was
published (Latin and old German of the early 18th century), this
material was not accessible to most zoologists, as a result of which
some disagreements arose in our day. Knowledge and iconography of
the species was poor.*

The morphological characteristics of the species have been pre-
sented in completely identical, though short, form as far as sufficiently
detailed descriptions of the series of taxa and forms. Geographic varia-
tion of species within the USSR has, with few exceptions, been revised
or critically reevaluated. Variation in those parts of geographic ranges
lying beyound the boundaries of the USSR follow the most recent
available data, but were not critically reviewed; the described forms
are listed and general notes provided.

All synonymys are re-examined and given in an extremely limited
scope according to the following principles: only species synonyms
are included. Generic synonyms are mentioned only if the type of the
genus is a species represented in our fauna, or if the generic reference
has significance in that it sometimes has been applied (or previously
applied), or may be applied to our species. All references are included
in species synonyms in the following cases: 1) if the name was ар-.
plied to a specimen of a species in our territories; 2) if the name was
applied to a specimen of a species taken elsewhere, but is significant
in that it is applied or has been applied but is now used for a Soviet
form; 3) if the name was given to a specimen from another territory
and not applied to our form, but may have significance for some

* Steller’s text is now known to be inaccurate in certain respects — Sci. Ed.

хи

Soviet forms; 4) the names applied to type species are always pre-
sented regardless of their type locality within or outside of the USSR.
Actual names in the species synonymies are not alloted, but are placed
in the accounts of subspecies inhabiting our country, and synonymies
are also given in relation to the particular subspecies. Synonymies of
subspecies distributed outside the USSR are not given. Synonymies of
categories above the genus are not mentioned, nor are references to
the descriptions and names of these higher orders which, if not stated
otherwise, are given by Simpson (1945).

Names applied to domestic forms, following the rationale and
usage of the first volume, are not used for wild species; rather the
name assigned the wild taxon is employed for the species as a whole.

The total number of mammalian species approximates 3500, and
the number of species of our fauna is about 300 (Heptner, 1956).

Each author’s contribution in the compilation of this volume was
as follows: V.G. Heptner wrote all the general characters of the major
taxa (order, family, genus), the entire sections concerning the sea cow
(including translation of Steller’s text) and the Afghan fox, and all
tabular material. He worked out the synonymy of all species and also
wrote the sections dealing with the “Description”, “Systematic Posi-
tion”, “Geographic Distribution”, and “Geographic Variation” for all
species described in this volume, and also prepared all maps of geo-
graphic range presented in the volume. P.R. Yurgenson wrote the
section on “Biology” and “Practical Significance” for the three spe-
cies of bears, all species of the mustelid family, and the American
raccoon. A.A. Sludskii wrote the same sections concerning the corsac
fox, jackal and red wolf [dhole] whereas the section on “Biology” and
“Practical Significance” for red fox and arctic fox were contributed by
A.F. Chirkova; N.P. Naumov wrote the same sections for the gray
wolf, and A.G. Bannikov, for the Ussuri raccoon (raccoon dog).
Authorship of the separate parts is indicated by bracketed initials at
the end of the corresponding sections. Integration of the [above] sys-
tem, species arrangement, their limits, number and limits of subspecies
as well as all the synonymies are the responsibility of V.G. Heptner.

The literature up to the period 1961—1962 was more or less com-
pletely employed. Some works which appeared after that period were
not referred to, especially the information provided by I.M. Gromov,
A.A. Gureev, G.A. Novikov, LI. Sokolov, P.P. Strelkov, and K.K.
СрарзКи (“Mammal fauna of USSR” 1963). We have succeeded in
only individual cases in making use of data from papers published

Xlil

after this period. Literature references are usually given parentheti-
cally by surname and year, initials being given only when more than
one author has the same surname. References to author’s unpublished
data, personal communications, or letters are given without reference
to year, but the author’s initials are given to distinguish them from
published citations.

The authors have, of course, included their unpublished personal
observations in this work, and this is sometimes referred to as noted
above. In addition much unpublished material of different types was
kindly supplied to the authors by G.F. Bromlei (Vladivostok), N.K.
Vereshchagin (Leningrad), N.N. Vorontsov (Moscow), S.V. Kirikov
(Moscow), V.V. Kozlov (Krasnoyarsk), A.P. Korneev (Kiev), V.P.
Kostin (Tashkent), B.A. Kuznetsov (Moscow), М.Г. Larina (Saratov)
Kh. I. Ling (Tartu), R.N. Meklenburtsev (Tashkent), L.G. Morozova
- Turova (Moscow), V.N. Nadeev (Irkutsk), T.A. Pavlenko (Tashkent),
V.Ya. Parovshchikov (deceased) (Arkhangel’sk), V.N. Pavlinin
(Sverdlovsk) V.A. Popov (Kazan), S.K. Priklonskii (Oka preserve),
В. Rausch (USA), С. Repenning (USA), М.М. Rukovskii (Moscow),
N.V. Rakov (Saransk), V.E. Sokolov (Moscow), V.N. Skalon (Alma-
Ata), S.U. Stroganov (deceased) (Novosibirsk), N.D. Sysoev
(Vladimir), K.A. Tatarinov (L’vov), V.P. Teplov (deceased) (Oka
preserve), V.V. Timofeev (Irkutsk), S.M. Uspenskii (Moscow), A.V.
Fedoseev (Bryansk), K.K. Chapskii (Leningrad), V.D. Shamykin (Mos-
cow), E.I. Shereshevskii (Moscow), E.I. Shcherbina (Ashkhabad), as
well as additional authors especially mentioned in the text.

Useful information concerning the distribution of the different
species was also kindly supplied by workers of all the district hunting
inspection centers. These were Starodubchenko (Kiev), Lebedev
(Volgograd), Mikhailov (Saratov), Zakharov (Murmansk), Gusev
(Rostov-on-Don), Khmelevskii (Orenburg), Sukhikh (Belgorod), Ivanov
(Yakutsk), Kuz’min and Fofanov (Novosibirsk), Bel’skii (Vladivostok),
Anashkin (Ulan-Ude), Samsonov (Blagoveshchensk), and Sysoev
(Khabarovsk).

Figures for this volume were assembled by V.G. Heptner. Animal
illustrations were prepared by our famous “animalists”, А.М. Komarov
and N.N. Kondakov. Line drawing of all of the animal species discribed
as well as five of the color plates, were made by A.N. Komarov.
Three plates—weasel, red fox and sea cow were done by N.N.
Kondakov. The painting of the Steller’s cow is considered a trial
reconstruction of the species—a collaboration between N.N. Kondakov
and V.G. Heptner. The drawing of tracks were, with few exceptions,

XIV

taken from Prof. A. N. Formozov’s books based on his original sketches
from nature.

The skull drawings were done by V.N. Lyakhov based on mate-
rials in the Zoological Institute of the Academy of Science, under the
supervision of Prof. G.A. Novikov, and were previously published in
his book (1956). The original drawings are in the possession of the
Zoological Institute of the Academy of Science; thanks to the courtesy
of Prof. O.A. Skarlato and Prof. A.A. Strelkov, the Institute has granted
permission for their use in this book.

Photographs and other illustrations related to the biological sec-
tions of the volume were contributed by various persons, but were
mainly taken from the archive of Prof. V.G. Heptner (especially those
of mustelid and bear families). A significant portion was provided by
A.A. Sludskii (Alma-Ata), Yu.K. Gorelov (Badkhyz preserve), S.V.
Marakov (Kirov) and A.G. Pankrat’ev (“Kedrovaya pad” preserve).
Those supplying other photographs are cited at appropriate places. All
photographs are original, and published here for the first time. Col-
leagues V.N. Orlov and I.P. Mitina of the Department of Vertebrate
Zoology at Moscow State University provided great assistance during
the preparation of this book.

We here once more acknowledge with thanks all the above-men-
tioned persons, organizations and others who helped in the production
of this book. The authors especially acknowledge Prof. V.A. Popov
(Kazan’) and Prof. N.I. Larina (Saratov) who provided great help in
the preparation and reading of the manuscript, and who made many
useful suggestions.

All work was to a great extent accomplished in the Zoological
Museum, Moscow State University, whose rich collections were used.
The authors express their gratitude to prior and present directors of
this Museum: Professors S.G. Soin and N.A. Gladkov.

The first volume “Mammals of the Soviet Union” was received in
our country, and in other countries as well, with great appreciation
and was officially and unofficially reviewed and put out in German
translation. The authors hope that this volume will also be acceptable,
even though they are quite sure that their work is not free from spe-
cific and general errors. The authors kindly request those who use
their book to inform them of any errors noticed and to send necessary
corrections to the following: Professor Vladimir Georgevich Heptner,
Zoological Museum, Moscow State University, Herzen St., No. 6,
Moscow K-9 [U.S.S.R.].

Table of Contents*

FOREWORD TO THE ENGLISH EDITION У
FOREWORD 1х
SYSTEMATICS OF THE CLASS MAMMALIA XIX
KEY FOR IDENTIFICATION OF ORDERS хх!
ORDER SIRENIA ILLIGER, 1811 1
Suborder Trichechiformes Hay, 1923 18
Family Hydrodamalidae Palmer, 1895 18
Genus Hydrodamalis Retzius, 1794** 18
Hydrodamalis gigas (Zimmermann, 1780) 18

(Steller Sea Cow)
Description and Biology 19
Geographic Distribution and History of Extinction 41
ORDER CARNIVORA BOWDICH, 1821 51
Key for Identification of Families in the Order Carnivora 64
Family CANIDAE Gray, 1821*** (Wolves and allies) 66
Key for Species Identification of the Wolf Family 74
Subfamily Caninae Gill, 1872 78
Genus Nyctereutes Temminck, 1839 (Raccoon Dog) 78
Nyctereutes procyonoides (Gray, 1843)**** 82
Diagnosis 82
Description 83
Systematic Position ; 87
Geographic Distribution 87

*Pages 1000—1001 in Russian original. The English table of contents is not a
literal translation.

**Palmer, 1895 in Russian original — Sci. Ed.

***Fisher, 1817 is now considered to have priority — Sci. Ed.

****1843 in Russian original—Sci. Ed.

XVI

Geographic Variation 91
Biology 92
Practical Significance 120
Genus Canis Linnaeus, 1758 (Wolves) 124
Canis aureus Linnaeus, 1758 (Jackal) Ae
Diagnosis 129
Description 129
Systematic Position 132
Geographic Distribution 134
Geographic Variation 140
Biology 141
Practical Significance 160
Canis lupus Linnaeus, 1758 (Gray Wolf) 164
Diagnosis 165
Description 166
Systematic Position 175
Geographic Distribution 175
Geographic Variation 182
Biology 198
Practical Significance 262
Genus Alopex Kaup, 1829 (Arctic Fox) 271
Alopex lagopus (Linneaus, 1758) (Arctic Fox) 274
Diagnosis PATS)
Description 275
Systematic Position 279
Geographic Distribution 279
Geographic Variation 287
Biology 292
Practical Significance 376
Genus Vulpes Oken, 1816* (Foxes) 385
Vulpes cana Blanford, 1877 (Afghan Fox) 390
Diagnosis 390
Description 390
Geographic Distribution 393
Geographic Variation 394
Biology 395
Practical Significance 395

*Frisch, 1775 is now considered to have priority — Sci. Ed.

XVil

Vulpes corsac (Linnaeus, 1768) (Corsac Fox) 396
Diagnosis BOT
Description 397
Geographic Distribution 402
Geographic Variation 406
Biology 409
Practical Significance 465

Vulpes vulpes (Linnaeus, 1758) (Red Fox) 469
Diagnosis 471
Description 472
Systematic Position 480
Geographic Distribution 482
Geographic Variation 485
Biology 504
Practical Significance 561

Subfamily Simocyoninae Zittel, 1893* 566
Genus Cuon Hodgson, 1838 (Red Wolf) 566

Cuon alpinus (Pallas, 1811) (Red Wolf) 571
Diagnosis ` 0971
Description 571
Systematic Position 574
Geographic Distribution 574
Geographic Variation 578
Biology 580
Practical Significance 586

Family Ursidae Gray, 1825** 586
Key for Species Identification of the Bear Family 596
Genus Ursus Linnaeus, 1758 (Bears) 597

Ursus (Ursus) arctos Linnaeus, 1758 (Brown Bear) 601
Diagnosis 604
Description 604
Systematic Position 612
Geographic Distribution 613
Geographic Variation _ 632
Biology 649
Practical Significance 676

*Dawkins, 1868 is now considered to have priority — Sci. Ed.
**Fischer, 1817 is now considered to have priority — Sci. Ed.

ХУШ

Ursus (Thalarctos) maritimus Phipps, 1774*

(Polar Bear) 678
Diagnosis 678
Description 679
Systematic Position 684
Geographic Distribution 685
Geographic Variation 694
Biology 700
Practical Significance MZ

Ursus (Selenarctos) thibetanus G. Cuvier, 1823

(White-chested, Black Bear) 713
Diagnosis 715
Description Ws
Systematic Position Gg
Geographic Distribution 720
Geographic Variation 723
Biology 724
Practical Significance 733

| LITERATURE CITED

{+ INDEX OF LATIN NAMES OF ТАХА**

*1774 in Russian original — Sci. Ed.

**The Russian original also included an Index of Russian Names of Taxa, omitted
here—Sci Ed.

+ These Sections appear at the end of part 1b—Gen. Ed.

Systematics of the Class Mammalia

9 The classification of major subdivisions (orders) adopted in the present
publication is based on the systematic monograph of the class by
Simpson (1945), who relied extensively on paleontological data. It is
very close to the system of classification proposed by M. Weber (1928),
which in its time was well founded on the anatomy of extant forms.
The orders proposed by Simpson. have long been widely recognized
and their grouping into higher taxa raises no serious objections.

Extreme division of the orders is a recent trend. Thus, marsupials
have been divided into three orders — polyprotodontids-Polyproto-
dontidia, caenolestids—Caenolestia, and diprotodontids-Diprotodontia,
cetaceans into two (toothless whales—Mystacoceti and toothed
whales—Odontoceti), primates into three (lemurs—Prosimiae, sim-
ians—Simiae, and tarsiers—Tarsioidea), and so on. All these proposals,
however, are not yet well founded. Moreover, among mammalian
systematists extreme division is not generally favored. Neither is it
commended here. An order should be primarily an integrating and not
a divisive concept. Otherwise the same situation will arise in mam-
malogy аз has already occurred in omithology, where the concept of
an order already has largely lost its significance; as a result the sci-
entific aspect of systematics of vertebrates has suffered greatly.

In the classification of orders under consideration here, only one
change has been made; 1.e., pinnipeds have been treated as an inde-
pendent order and not as a suborder of Carnivora. Based on the
systematics of recent mammals, these two groups separate naturally—
they are not less isolated than some other orders.

Based on already established views, Lagomorpha are treated here
as an independent order (Weber considers them only a suborder of
rodents). Hence, instead of the 18 orders of Simpson, we propose 19.
Of these, 10 (52.6%) are represented in our fauna, one—the sea cow—
is now extinct (Steller’s sea cow).

The main subdivisions within the orders in most cases adhere to
Simpson’s scheme. By and large his classification is the most well

founded. Nevertheless, there are a series of deviations, some signifi-
cant, mainly in interpretation of the content of the genus, which is
understood more broadly than is generally the case. In our opinion
this conforms better to the structure and principles of natural classi-
fication.

The systematic classification adopted is given in the chart below.
Orders present in the USSR fauna are marked with an asterisk. (V.H.)

CLASS MAMMALIA

Subclass PROTOTHERIA Order
MONOTREMATA
Subclass THERIA
Infraclass METATHERIA Order MARSUPIALIA
Infraclass EUTHERIA
* Order INSECTIVORA
Order DERMOPTERA
* Order CHIROPTERA
Cohort UNGUICULATA Order PRIMATES
Order EDENTATA
Order PHOLIDOTA

ic Order LAGOMORPHA

Cohort GLIRES * Order RODENTIA

Cohort MUTICA * Order CETACEA

Superorder FERAE | * Order CARNIVORA
* Order PINNIPEDIA

Superorder Order
PROTUNGULATA TUBULIDENTATA

Cohort Order PROBOSCIDEA
FERUNGULATA| Superorder Order HYRACOIDEA
PAENUNGULATA | * Order SIRENIA
Superorder * Order
MESAXONIA PERISSODACTYLA
Superorder * Order

PARAXONIA ARTIODACTYLA

1).
2
3 (4).
4 (3).
5 (6).
6 (5).
7 (10).
8 (9).
9 (8).

10 (7).
11 (14).

РЗ)
1312).

14 (11).

Key for Identification of Orders

Hind limbs absent; fishlike body with large bilobate caudal
fins Зенона У СЕТАСЕА.
Hind limbs present; body not fishlike and tail, if present,
not in the form of a bilobate fin.

Forelimbs in the form of leathery wings............

Forelimbs of a different structure.
Fore- and hind limbs very short and resemble paddles,
1.e., all the digits entirely enclosed in a common

IMLS OUI EM Cesc ei ore ako eit tal eee PINNIPEDIA.
Fore- and hind limbs of a different structure, not paddle-
shaped.

Feet possess hooves.

Only one hoof on each limb....... PERISSODACTYLA.
On each limb, two large hooves and two small ones above
RSI een о АИ ae ARTIODACTYLA:!

Hooves absent (claws present).

Between large chisel-shaped incisors and molars on jaws,
a large toothless gap (diastema) occurs; its length not less
than length of entire row of molars on corresponding jaw.
Canines absent.

TwoAncisors'on upper jaws... : + lense RODENTIA.
Four incisors on upper jaw; behind each large sharp incisor
is located a minute blunt tooth... .... LAGOMORPHA.

Diastema between incisors and molars absent or much
smaller than length of molar row. Canines present.

'Limbs differ in structure in camels. The leg terminates in a broadened callused
pad and true hooves are absent. There are two greatly broadened nails. Camels have
not been included in this table as they are domesticated animals (wild camels are

extinct).*

*It is now believed that wild populations of Bactrian camels still survive in China
and Mongolia—Sci. Ed.

XXil

15 (16). Anterior portion of muzzle extended into well-developed
small proboscis. Anteriormost tooth on each jaw, or only
upper jaw, much larger than adjacent tooth. INSECTIVORA.

16 (15). Anterior portion of muzzle does not form a proboscis.
Anteriormost tooth on each jaw not larger than one next to
О. ere A, и a, CARNIVORA.’

*If the structure of teeth is different, proboscis is always present and furthermore
the front leg extremely short, feet very broad with huge claws, and set on edge with
inner surface facing backward (moles).

3On the coast of the Commander Islands, skulls of the extinct Stteller’s cow—
a member of the order of sea cows (Sirenia}—have been found. They are distinguished
by upper and lower jaws measuring 60 cm in length, in which teeth or traces of them
(alveoli) are absent (fig. 6).

ORDER OF SEA COWS,
OR SIRENIANS

Ordo Sirenia Illiger, 1811

Superorder of Probosciforms*

Superordo PAENUNGULATA Simpson,
1945

Order of Sea Cows, or Sirenians
Ordo SIRENIA ППоег, 1811**

Mammals of large size, entirely monomorphic,*** highly special-
ized as typical aquatic animals. The species of this order represent one
of the extreme degrees of specialization among mammals.

Body elongated, fusiform, and in general appearance, more similar
to that of large dolphins, but as compared to the latter, and in relation
to total animal size, the head is smaller and located on a short, movable
neck not noticeable externally in all animals. In connection with the
low mobility of the animal, the general body constitution is much
heavier than that of dolphins and the body diameter in its thickest part
is much larger. There are no external hind limbs. The hind part of the
body is elongated and thins gradually, rather than abruptly, into a
relatively thick tail region which turns into a wide, flattened horizon-
tally oriented tail fluke, lacking internal skeleton (other than the vertebral
column and the connective tissue). The posterior edge of the fluke
forms a convex arch and the whole fluke is blade-like (Family
Trichechidae—manatees), or else the posterior edge is deeply incised
in the middle and the tail is bilobed (remaining families).

The anterior extremities are in the form of true flippers, without
any [external] signs of differentiation into digits. Some species have
nail rudiments on the П-У fingers (several manatees). The flippers are
freely mobile in the shoulder joint and (in contrast to cetaceans) pos-
sess a considerable mobility in the elbow and hand articulations (see
p. 30). The animal leans upon them, raises the anterior part of its body,

*In Russian original, slonoobraznykh; literally, “of elephant-like form’—Sci. Ed.
**Co-edited by Daryl Domning (D.D.), Howard University, Washington, D.C.
***Except Dugong — D.D.

moves, and performs other relatively complicated movements (arrang-
ing food in its mouth, etc.).

On the head, in connection with the peculiar structure of the skull
(see below), the upper lip is extremely developed, forming with the
nasal region a massive fleshy mass hanging down over the lower,
normally developed lip, and represents a sort of wide, blunt and short
proboscis. The mouth opening lies on the lower surface of the head.
The shape of the “proboscis” varies in different forms (higher or wider),
but in all forms it is highly movable, capable of changing its shape, and
plays an important role in collecting food. In some animals (for ex-
ample the manatees, Trichechus), the actively movable lateral parts
can even mimic to a degree, a prehensile organ. It acts like the index
finger and thumb of humans, and by bending it to the left and to the
right the animal can grasp plants. The nostrils are located on the an-
terior upper part of the “proboscis”; they are crescent-shaped and closed
(open only during inhalation and exhalation). The eyes are small and
round; eyelids are devoid of eyelashes but richly supplied with special
glands, secreting abundant viscous fluid that protects the eyes from the
influence of water.' The Harderian gland is highly developed but, in
contrast to cetaceans, the lachrimal glands are apparently absent. A
sclera is not formed.* A well developed and highly movable nictitating
membrane is present. The ears are located far behind the eyes and are
represented by very small orifices; a pinna is completely absent.

The hair cover of the body is completely, or almost completely,
absent and is represented by fine hairs scattered singly on the body, and
having the structure and, apparently, function of sensory organs. This
fulfills the need for high tactile sensibility of the skin, important for an
animal living among aquatic vegetation in coastal zones. In the fetus,
the number of these hairs is considerably greater. On the upper lip and
partway around the mouth are found a large number of hairs; usually,
however, they are short and do not form the “whiskers” characteristic
of some pinnipeds. The majority of the hairs are elastic and bristle-like,
but some are softer. Some hairs have a tactile function while others,
especially the coarser ones, are considered a device for assisting the
grasping of food with the upper lip.

'In animals removed from the water, this fluid accumulates in the corner of the
eye and then flows out. The legend of the “weeping sirens” has originated from this,
and so does the name lamantin (from the French Lamenter: to weep). This oily liquid
is, apparently, a product of Meibomian and Harderian glands.

*Meaning not clear—Sci. Ed.

In the majority of species, the skin has a very thick connective
tissue layer (about 20—25 mm) and very thin epidermal layer (cuticle
is about 0.1 of the total thickness of the skin). The skin [in manatees]
is thicker on the sides than on the back as is characteristic of mammals
in general; on the abdomen, it is approximately half as thin as that on
the sides, but on the back, it is thicker than on the abdomen. Sometimes
(Steller’s sea cow, see below), the epidermal layer is especially devel-
oped and strong. In some species (Amazon manatee, Trichechus
inunguis) the border of the caudal fin bears а hard comification border
(or is included in it) which may be detached and again re-formed.’ The
fatty layer is well developed.

The [skin] color is dark, more or less evenly brown, brownish or
black in color; from below, it is lighter in some animals, and sometimes
of a rare-meat color.

There are two mammary glands.

The female genital opening is slit-like, placed fairly far in front of
the anal opening. The testicles lie in the abdominal cavity (far anterior,
near the diaphragm); the male copulatory organ is long with highly
developed corpora cavernosa, lies hidden under the skin, and contains
no bony structures.

The skulls of all species of this order are completely monomor-
phic,* very peculiar and differ greatly in general structure from those
of other mammals. They have some similarities with the skull of artio-
dactyls and perissodactyls, and particularly with those of elephants and
hyraxes, but they sharply differ, in general and in details, from the
skulls of cetaceans.

In its general features, the skull is characterized by the fact that the
nasal openings are greatly shifted and directed upwards, and the pre-
maxillae are large in size. In some animals they are very large and
massive and are bent downwards in their anterior half (symphysial
part), taking on a beak-like form, and giving the skull a curved form
in side view (Hydrodamalidae, Steller’s sea cow; and especially
Dugongidae, dugong). In other forms, the premaxillae are less massive
and extend forward, but are not, or are only slightly bent downwards
(Trichechidae), and do not fuse together in a symphysis.** The nasal
processes of the premaxillae extend far backwards, reaching the nasal
bones. Together with the enlarged maxillary bones, they delimit the

*Judging by some figures of Steller’s sea cow (see below), they apparently
possessed such a structure on the posterior border of the fin.

*Except Dugong—D.D.

**Not true; symphysis is formed—D.D.

nasal cavity. The nasal bones are very small and shifted far posterior,
forming [part of] the rear wall of the wide nasal opening, on the bottom
of which the widely opened vomer is found. The nasal bones are some-
times absolutely rudimentary [or absent]. The maxillae are large, and
the palate is long.

The supraoccipital is large, greatly extending forward, but not
wedged between the parietals. The suture between the parietals disap-
pears at an early stage and a single parietal is formed, constituting the
cranial roof. The frontals are separated and with large supraorbital
processes. The squamosal bone has a large zygomatic process, to which
the jugal bone is connected. The space between the exoccipital and
squamosal is only partially filled by the petrosal bone (petrosum); [the
remaining space] consists of a large opening (filled with connective
tissue in the living animal) leading to the cranial cavity. The tympanic
bone (tympanicum) is quite closely connected [fused] to the petrosal
bone, forming the petrotympanic bone (petrotympanicum). The tym-
panic bone is in the form of a fairly massive half-circle, but it does not
form a bony auditory capsule (bulla). The petrotympanic bone is not
fused with the skull but freely articulates and connects with the squa-
mosal bone. The anterior and posterior lacerate foramina (for. lacera)
are fused in a common large one. The lachrymal bone is small, not
perforated, and in some animals it is connected with the surrounding
bones. There is no lachrymal canal. The palate 15 long and the palatine
bones, being small and shifted posterior, play a very insignificant role
in its composition.

The mandibular bones are massive and heavy, connected in an
elongate symphysial portion; they form a single bone. The anterior part
of the lower jaw is recurved downwards, and the degree of its bending
corresponds to the form and position of the premaxillae (most strongly
in the dugong, and to a somewhat lesser degree in Steller’s sea cow).
The ascending ramus of the lower jaw has strong, well developed
articular and coronoid processes, directed forward and of various rela-
tive size in various forms, and a massive broad angular process. There
are no air cavities in the skull.

The dental system is highly peculiar and differs greatly from that
of other mammals. The cheek teeth [in Trichechus]* are absolutely
identical with a short crown carrying two transverse crests, each of
which is formed from three tuberosities. During occlusional wear, the
masticatory surface is flattened. The number of simultaneously func-
tioning teeth in one jaw ramus of a mature animal is 3—4 or even 2

*D.D.

(Dugong) and from 5—6 to 8 (manatees, Trichechus). All the teeth are
considered as true molars [some are deciduous premolars]', and inci-
sors, premolars and canines are absent. Only in male [and a few old
female]* dugong one permanent incisor erupts on each side of the upper
jaw, converted into a peculiar small (up to 10 cm) “tusk”, directed
downward.

With wear, the teeth [in Trichechus]' are replaced by new ones, but
not by the old tooth being pushed out by the new one growing under it.
The newly formed teeth appear at the extreme rear of the toothrows and
at the same time the anterior ones (“‘old” teeth) fall out, the crowns having
almost worn out and lost their roots, which are well developed in “younger”
teeth. Accordingly, the toothrow is continuously shifted forward. This
takes place because the posterior side of the bony septum between teeth
(i.e. the septum between each two successive teeth) is resorbed, while
on its anterior surface (i.e. adhering to the rear surface of the tooth in
front) this bony septum grows forward. In manatees, replacement of
teeth occurs throughout the whole life and the number of molars is, as
a matter of fact, not limited. In manatees, for example, in every jaw
quadrant (“half’ jaw) not less than 30 teeth or possibly more? are
replaced throughout the animal’s life. If we take into consideration only
the teeth functioning at a given moment, the dental formula for a fully

mature dugong (Dugong) is IoC“ POM = 10, while in the case of

ee 1.0 3
young animals of this species, it is СОРМ = 28*.

The biological significance of this unusual method of tooth replace-
ment in manatees probably lies in the fact that in feeding on aquatic
vegetation much sand** is obtained and the teeth are worn out rela-
tively quickly.

*D.D.

'This manner of tooth replacement -from behind forwards occurs only in
elephants.*** However, the number of teeth replaced in the course of life does not
exceed 6 and only one tooth is functioning in each jaw ramus (in all 4 cheek teeth).
Sirenians and elephants are also similar in that the tusks in both are formed by the
incisors. The transverse orientation of the crests on the teeth in sirenians is similar
to the corresponding structure in primitive Proboscidea.

*In Russian original, «7c 2 p2® vie 26” which is erroneous—D.D.
103

**Probably siliceous phytoliths in the vegetation—D.D.
*** And some macropod marsupials—D.D.

Overall, a reduction in the dental system occurs in this order, and
in one species (Steller’s sea cow, Hydrodamalis) teeth are lost com-
pletely.

This reduction is connected with, and probably due to, the fact that
in all species, in both the upper and lower jaws there are large elon-
gated horny plates occluding with each other in front of the tooth rows.
On the lower jaw, the plate is located on the symphysial part which 1$
bent downward, while on the upper jaw, the plate lies on the lower
concave surface of its anterior part. Such a system 1$ used for seizing
and grinding food and has no analogue among other mammals. The
small cornified part lying in the region of the anterior end of the
premaxilla in even-toed ungulates has, however, some functional and
morphological similarities to the upper plate in sirenians (see vol. 1).

Owing to the great reduction of the skeleton of the posterior ex-
tremities, true sacral vertebrae are absent,* and not differentiated from
the lumbar vertebrae; the caudal region is long. In the genus Trichechus,
the number of cervical vertebrae is six.* The number of trunk vertebrae
in various species is 19—24; caudal—22-29, and 35 (Hydrodamalis,
Steller’s sea cow); ribs, 14—19 pairs.** Thoracic vertebrae are very

*True of manatees, but not dugongs — D.D.

‘The number of the cervical vertebrae in other species cannot apparently be
considered established. According to Weber (1928) there are 7, while Mohr (1957)
stated that in the dugong and Steller’s sea cow there are “?6 ?7”. Steller (1745—
1753), who gave a detailed description of the Steller’s sea cow anatomy, noted 6
cervical vertebrae.

The skeleton in the Zoological Museum of Moscow University has six vertebrae
(V.G. Heptner), but it is possible that one is missing. In the skeleton found in the
Zoological Museum of the Academy of Science (Leningrad) are seven cervical vertebrae
(K.K. Chapskii), but it cannot be excluded that long ago in assembling the skeleton,
some vertebrae might have been added to reach the number normal for mammals. A
skeleton present in the Museum of Kiev University has seven vertebrae, but it is possibly
more correct to consider them six; one thoracic vertebra may be placed with the cervical
ones (A.P. Korneev). It is unlikely, however, that Steller could commit such a terrible
mistake, and therefore it subsequently was agreed to consider the number of the cervical
vertebrae in this animal as being six until further special investigation is carried out.

A solution of this problem based on museum skeletons is at the present time
difficult, because all of them are actually composed of bones from different individu-
als. Only Evermann (1893) bought in 1891 an almost complete skeleton from the
Commander [Islands] which was found “а short time” before that. The finder con-
firmed that the skeleton was found not far from the water’s edge under a few inches
of sand. The skeleton contained the complete set of cervical vertebrae which, after its
study in the National Museum of Washington, was shown to be 7.

The deviation in the number of cervical vertebrae is a rare exception among
mammals. Besides manatees, only the two-toed sloth, Choloepus hoffmanni, has 6
vertebrae, and the three-toed sloth, of the genus Bradypus, has 9 vertebrae.

** 14—19 pairs among living species — D.D.

19 few and only 2-3 or 5 of the anterior ribs (Steller’s sea cow) are
connected to them, while the remaining ribs are free. Clavicles are
absent.

The scapula is well developed and carries a normal spine. The
articulation of humerus with scapula is quite free, the elbow joint is
freely movable and the wrist joint has a considerable degree of mobil-
ity. Ulna and radius are separate and only in a few forms do they fuse
distally in older animals.* In some young animals the whole set of
carpal bones 1$ present, but in mature ones, they fuse together into two
or three large bones. The manus 15 pentadactyl and of normal construc-
tion; polydactyly is not found, and supernumerary phalanges do not
normally occur in the species. Little is known, in contrast to cetaceans,
of the structure of the forelimb skeleton in limiting their capability for
different movements, such as acute bending of the hand in conveying
food to the mouth, etc.

The skeleton of the hind limbs is utterly lost and only in a few
species is there a totally insignificant rudiment of femur with an articu-
lating head. The pelvis is represented by only the ischium and ilium,
which are united together [in dugongids] into a rod-shaped bone in
which its constituent elements are not well differentiated. There is no
acetabulum.

The bones of the skeleton have no marrow cavities, and all of them,
including those of the skull, are characterized by what is termed
pachyosteosclerosis—Haversian canals are closed, all bone tissue is
very compact resembling that of elephant bones, and shows conchoidal
fracture. Moreover, the bones are relatively massive. All these features
make the skeleton very heavy. Apparently, these features are of adap-
tive significance for animals living in coastal waters, bringing their
specific gravity close to that of water, thus facilitating underwater
submergence, etc.°

The kidneys are lobular as in true marine animals (whales, pinni-
peds), the intestine is very long and exceeds the body length by 13—20
times, the large intestine is very long, only a bit shorter than the small
intestine, and sometimes longer (Steller’s sea cow). The caecum is well
developed and in some species has accessory diverticula. The stomach
is a complex organ, with a sharp constriction between the cardiac and
pyloric parts, an additional blind sac or outgrowth in the cardiac region,
and two blind sacs in the pyloric part.**

*Not true; in most they fuse both proximally and distally — D.D.

‘Paleontological data show that pachyostosis appears in the evolutionary
sequence during the transition to life in water.

** Actually part of duodenum — D.D.

The uterus is bicornuate and the placenta zonal.

With the exception of some differences in size and development of
tusks in males [and females] of one species (Dugong), there is no
sexual dimorphism.

The body length of smaller species reaches 200—300 and may
reach 360 cm. Body diameter, in its thickest part, is about 150—155 cm
(for a length of 203—247 cm) and the weight is up to 400—600 kg. In
the largest species (Steller’s sea cow), the body length reaches 800 cm,
the diameter of the trunk about 620 cm and the weight up to 3200 (200
poods) and 4000 kg (morphological structure according to Weber, 1927,
1928; Mohr, 1957 and other references, and material in the ZMMU
[Zoological Museum, Moscow University]).

They are specialized aquatic animals which do not come out on
land,° and are associated with shallows of coastal rivers, in particular
with calm bays, lagoons, and river mouths, and with rivers rich in
aquatic vegetation. They do not live far from shores, and particularly
[do not inhabit] pelagic parts of the sea. Three species are purely
marine, one marine and freshwater,* and one is strictly freshwater.
Although most are inhabitants of tropical, warm water, one species
inhabits the cold temperate zone. They are herbivorous forms, feeding
on submerged and floating aquatic plants (“grazing”) along the shores.
They are very sluggish, flaccid, and clumsy animals [on land] of quite
low mental level. They are usually found in small groups. Young are
usually single.

This order is widely distributed, the major part of its range being
characteristic of littoral (coastal) marine animals and only in part char-
acteristic of the ranges of animals associated mainly with the lower
courses of larger rivers. Therefore, everywhere the range is typically
linear (Heptner, 1936). The general range is divided into four isolated
parts, differing greatly in their size—two lie in the Atlantic and two in
Indian and Pacific oceans. These parts are not continuous, but divided
into smaller and larger sections.

In America, the range (reconstructed, including transgressions in
the north) extends to the southern coast of North Carolina (Wilmington)**

Га sea cow is exposed on land or subjected to drying in some other way (for
example, ebb-tide), it does not die of this, but as do whales, is “crushed” by its own
weight and is overheated.*** However, these animals cannot move themselves on
land.

*Two considered marine, and two both marine and freshwater—D.D.

**To New England—D.D.

***Мо evidence for this—D.D.

‘rauday ‘N'A ‘эидзэреа JO 35609 OY) UO ("ПИ VYOsnp Suosnq) Buosnp sy) JO р1оЭэл [емащецхо ие $1 BIS
ивэцецэирэи{ э4з 30 wed wsojseo эф ul julod oy] ‘(зривз| оэцэлеэзеу ‘воцуу YyINog) keep прапор 10 ‘Ajsnotasid 35691 зе
‘Q0UdLINDIO 91415504 зпа ‘иоцеиноуит JO ээиэзае OY} эзеотриг ззлеш иопзэпо ‘ешэнс ‘SUBIUDIIS JO SMOD BAS JO 1эрло dy) JO эвивл рэзопцзиозэч ‘1 ‘8 15

> a
ИА М

an
:

y
=

and occupies the coasts of the Gulf of Mexico, islands of the West
Indies, and Central America, as well as part of the northern and north-
eastern coast of South America—the shores and river mouths in
Colombia, Venezuela and Guiana to Cape Norte in the east (slightly
southeast of Maraca Island, Brazil), lying somewhat north of the mouth
of the Amazon. The range of this [partly] marine species of the West
Indies (Trichechus manatus) does not, apparently, reach the mouth of
this river.’ Moreover, the range in South America (of the freshwater
species 7. inunguis) covers the basin of Orinoco* and Amazon basin
far to the west including, in any event, the region of a right-hand
tributary of the Amazon, the Rio Madeira (according to the data of
Hatt, 1934; G. Allen, 1942; Cabrera, 1960).’

At the present time the range is strongly modified and animals have
been exterminated in most places, and north along the Atlantic coast,
manatees are no longer found beyond the southern extremes of Florida,*
and where they are protected.

In West Africa, the range occupies coastal lagoons and river mouths
from the Senegal River on the north to the Cuanza River in northern
Angola. The species inhabiting this region (7. senegalensis) lives in
fresh [and salt] water and they penetrate upstream in the majority of
West African rivers, especially the large ones. Thus, in the Niger, their
range includes its left tributary, the Benue, but in the Niger itself, the
range extends almost to Timbuktu. Manatees are widely distributed in
the Congo basin.** Their upstream distribution along rivers is generally
restricted by waterfalls and rapids. Reports and suggestions concerning
the occurrence of manatees in Lake Chad and its basin are mistakes

’The range suggested by Cockrum (1962) is, apparently, not true even for the past.
There are direct indications that owing to hydrological conditions (Murphy, 1936 and
others), manatees did not occur to the east beyond the place mentioned, and were not
found on the coast of northern Brazil east of the mouth of Amazon River. They did not
reach Cape Sao Roque (the eastern point of the continent) and have not spread to the
south, although, in the area southward from the Cape, along the eastern coast, the
conditions for them were apparently favorable.*** Cockrum (1962) assumed that the
area of distribution of these animals extended along the coast of South America to the
mouth of the La Plata [River].

*“Rio Madevia” and “Вю Madeva” given by Mohr (1957) are probably misprints
instead of “Rio Madeira” (type locality for this species).

*Not true—D.D.

**No evidence for this statement—D.D.:

***West Indian manatees occur in this region today—D.D.

lacking foundation,* as are data about the presence of manatees in the
Great [Rift] Lakes of Africa.?

In the Indian and southwestern parts of the Pacific Ocean, the range
of the dugong (Dugong dugon) occupies the greatly extended shoreline
from East Africa and the Red Sea in the west to eastern Australia, the
Solomon and Marshall islands, and the Ryukyu islands. The range in the
above-mentioned areas was probably discontinuous from time immemo-
rial, but because of inadequate information it is now impossible to
reconstruct the true range. The range is divided at present into separate
parts.'? Along the eastern African coast, the range extends in the south
to Delagoa Bay at 26° $. Lat., a little south of Lourengo Marques, but
in earlier times it may have been extended to the south (to the Cape of
Good Hope?), and occupied the whole Red Sea along the southern coast
of the Arabian peninsula, though not east of Aden. It occupies the coast
of Madagascar, mainly the west and the small islands off the African
coast (Mafia** and others).'!

Yet, the dugong is apparently absent and probably has not been
present for a long time in the majority of places in the north of the
Indian Ocean; i.e. the southern coast of the Arabian peninsula, in the
Persian, Oman, and Bengal gulfs, and the shores of Indostan.*** The
range here (the second part of the Indo-Pacific Ocean range) occupies
only the most southerly part of the west coast of India, starting from
Cananora (Canara [Cannanore], about 12° N. Lat.) southward and to
the southernmost part of the eastern coast of the peninsula lying adja-
cent to Ceylon, which is included in the range.

The third part of the Indo-Pacific Ocean region occupied by sea
cows is connected with extreme southeastern Asia, Australia and the
islands lying between them. In the south, the range includes the western
coast of Australia to the north of Shark Bay (about 25° S. Lat., a little

*There is now some evidence that manatees occur in the Lake Chad basin—D.D.

*Information concerning the occurrence of manatees in the past (about 1800) at
Saint Helena Island in the Atlantic Ocean, which in its time provided material for
zoogeographical speculation, is false and is probably based on the appearance of
elephant seals (Hatt, 1934; Murphy, 1936).

The range described here differs considerably from that presented by Cockrum
(1962, map) which was incorrect at least in part. For example, the occurrence on the
coast of Madagascar was not given although a series of reports were available.

**Off coast of Tanzania—Sci. Ed.

''Conceming the Mascarene, Comoro and other islands of the Indian Ocean and the
Mozambique Strait associated with Madagascar, there are no reports. Only Cockrum
(1962) includes the Mascarene islands in the range.

*** Archaic; Indian subcontinent—Sci. Ed.

south of Carnarvon), the northern coast, and the eastern coast south to
Botany Bay (near Sydney). In the west, the range covers the Greater
and Lesser Sunda islands. In the east, it covers the Solomon and Marshall
islands (there are no definite records from other islands of Micronesia
or Melanesia, in particular the Bismarck archipelago). In the north, the
range includes Sumatra, tle coasts of Malacca Strait (no definite records
are known from the Nicobar and Andaman islands), the coast of the
Gulf of Siam, the whole of the Philippine archipelago as far north as the
Batan islands, Taiwan and the Ryukyu archipelago north to Amami-o-
shima (Amamioshima, 28° N. Lat.). Thus the range includes the southern
coast and western extremes of New Guinea (concerning the northern
coast, records are lacking), and the Molucca Island group. In this re-
gion, the range occupies the Coral, Arafura, Timor, Banda, Java, Flores,
and Celebes seas, Macassar Strait, part of the South China Sea, 1.е. all
the internal seas lying between Asia and Australia, part of the East
China Sea, the extreme eastern coast of the Indian Ocean, the extreme
west part of the Pacific Ocean, and that part in the Micronesian region
(Blanford, 1876; Allen, 1942; Carter, Hill, Tate,* 1946; Cockrum, 1962).

A distinct, very small part of the range lies separated from other
parts in the Commander Islands [Hydrodamalis only].

A known event is the passage of a dugong through the Suez Canal
to the Palestinian coast (Aharoni, 1930).

Remains of Tertiary sirenians (fam. Dugongidae) are known far
from the present limits of the range—in Africa, Europe and even in
Jamaica.

Sea cows, as was previously mentioned, are considered as a sharply
characterized group, and thus, accepted by all modern investigators as
an order. However, a short time ago, the sea cows were sometimes
considered as a suborder (Sirenia) of the Order Subungulata, which,
besides Sirenia, includes Hyracoidea, Proboscidea and the extinct
Embrithopoda. Systematic and phylogenetic connections between all
three contemporary groups is unquestioned—as was reflected in their
combination in the Superorder Paenungulata (see р. хх). However, dif-
ferences between them, even by evaluating all palaeontological data, are
those normally recognized for ordinal rank. The differences between
Sirenia and both Proboscidea and Hyracoidea are undoubtedly greater
than those between Pinnipedia and Carnivora.

The systematic position of this order is, according to recent data,
determined by their previously mentioned relation to Paenungulata. This

*Misspelled “Teg” in Russian original—Sci. Ed.

order has, apparently, more common features with the order of elephants
and they can be considered as sister groups although the relation be-
tween them is quite distant. The nature of tooth replacement and some
of its peculiarities, which is not elsewhere found among mammals, the
transformation of incisors into tusks, some features of skull appearance
and others, are among these common characters.* The similarity some-
what increases in the comparison of the more primitive ancient forms of
both groups. Sea cows are sometimes considered closer to Hyracoidea.
In any case, there is no doubt (and this is very obvious) that the sea
cows have no relation either to pinnipeds or to whales. All the features
common with whales are no more than convergences, and close study
reveals deep principal differences between them. The greatest similarity
with whales lies only in the general appearance, the absence of the rear
extremities, and in some details characteristic of aquatic mammals in
general.

In the eighteenth century, at the beginning of the last century and
somewhat later, even competent authors (Pallas, for example) included
the sea cows in the order Cetacea (whales) as a special group (“grass-
eating whales’). However, even at that time, there were some indications
not only of the independence of sea cows as an order, but also of their
relatively closer relation to elephants. This point of view was, a long
time ago, generally accepted, though it must be emphasized that the
systematic isolation of this order is very sharp and that its relation to
elephants is very distant. The same also applies to ungulates, though in
the system of orders of the class, sea cows are always put between
elephants (or hyraxes) and one of the orders of ungulates [see p. xx, this
volume].

The order comprises two suborders: Trichechiformes, in which to-
gether with the extinct ones, all living forms are included, and
Desmostyliformes [now an order], including only extinct forms (one
family with perhaps five genera). The systematic position of this group
is, however, not completely clear, and some taxonomists consider it a
separate order, Desmostylia, which is close to Sirenia.

The suborder Trichechiformes comprises five families, of which two
(Prorastomidae — one genus, Protosirenidae — one genus) are extinct and
three living: manatees, Trichechidae; dugongs, Dugongidae and kapustniks,
or Steller’s sea cows, Hydrodamalidae.** The family Trichechidae is,

*In part, no longer considered true — D.D.
**Now reduced to a subfamily of Dugongidae — D.D.

apparently, considered the least specialized among the existing ones. Spe-
cies of this family have a relatively less developed, only slightly depressed
anterior (“ргетахШа”) part of skull [rostrum], complete dental system, and
6 cervical vertebrae. The tail is blade-like and single-lobed. This group
includes one genus, Trichechus. The family Dugongidae comprises four
subfamilies: Halitheriinae (7 genera), Miosireninae (1 genus) and
Rytiodontinae (one genus) are extinct, and the Recent Dugonginae with 1
genus Dugong. This last family is characterized by the considerable reduc-
tion of cheek teeth (to 2), the presence of tusks in the upper jaw of males,
the extremely enlarged and sharply depressed upper jaw, and 7 cervical
vertebrae. The tail is bilobed. The family Hydrodamalidae, kapustniks or
Steller’s sea cows, with | [recently extinct] genus, Hydrodamalis, is char-
acterized by the complete loss of teeth, a fairly highly developed and
depressed rostrum, and 6* cervical vertebrae. The tail is bilobed.'*

Evidence on the origin of the group is given by representatives of
the above-mentioned extinct families existing only in the Eocene, and
the family Dugongidae which is represented by its extinct subfamilies
in the Eocene—beginning especially from the Middle Eocene to Plio-
cene. The other existing families have no extinct representatives, or only
their Pleistocene remains are known (manatees).

The oldest Middle Eocene forms, such as Eotheroides, though show-
ing some essential differences from the existing ones (complete dental
system, apparently, since they were characterized by the absence of
horny plates, a fairly well developed pelvis, etc.), were true sea cows.
They are well linked with the existing forms by means of a succession
of forms. In the Middle and Upper Eocene, they were already very
widely distributed and, therefore, the origin of this order must be related
to the Paleocene. As was shown, the similarity in the structure of skull
and teeth of the more ancient sea cows with primitive proboscideans and
hyraxes is more evident than in the existing sea cows. The ancestors of

*Now known to be 7—D.D.

The above-mentioned classification of the order differs from that adopted by
Simpson (1945), mainly in the acceptance of Hydrodamalidae as a family (which he
considers as only a subfamily of Dugongidae). This is partially determined by the fact
that Steller’s cows possess 6 cervical vertebrae and not 7.* Other authors consider this
group also as a family (Weber, 1928; Mohr, 1957) even when there was the assumption
of 7 vertebrae. The order of the families as regards increasing specialization is also
changed. In sum of characters, the family Hydrodamalidae is closer to Dugongidae than
to Trichechidae and is considered as a further development of this type. The difference
in the number of cervical vertebrae sharply separates the families from one another.

A more thorough analysis of the extinct and existing forms and their relationship
is given by Simpson (1932).

sea cows must, probably, be searched for among land forms, especially
those which gave rise to the other Paenungulata, i.e. among the hyraxes
(order Hyracoidea) and proboscideans (Proboscidea) as well as ungu-
lates (Artiodactyla and Perissodactyla). However, direct “basal forms”
of the order are not known.

The existing species in the order are 5. In the genus Trichechus L.,
1758 (= Manatus Briinn.) there are 3 species: T. manatus L. (shores of
the Americas—see above), Т. inunguis Natt. (basins of Amazon and
Orinoco*) and Т. senegalensis Link (sea coasts and the lower reaches
of the rivers of West Africa). In the genus Dugong Lac., 1799
(= Halicore Ш.) there is 1 species—D. dugon Erxl.** (the coastal
region of the Indian Ocean and the adjacent southwestern Pacific). The
kapustniks, or Steller’s sea cows, Hydrodamalis Retzius, 1794—1
species—H. gigas Zimm. (coastal waters of the Commander Islands).

The greater number оЁ зресиез are endemic to the Atlantic Ocean,
with two associated with its western coast and one with its eastern
coast. One species is endemic to the Indian Ocean and small parts of
the Pacific contiguous to it. The practical value of the group is insig-
nificant. They are only hunted locally for their meat. In many places
these animals have been extirpated and their number is limited almost
everywhere. One species was completely exterminated (Steller’s sea
cow).

The USSR fauna comprises only | species, the kapustnik or Steller’s
sea cow, Hydrodamalis gigas Zimm., 1780—which constitutes 20% of
the species of the order and about 0.3% of the number of species of our
fauna (V.H.).

*Not found in Orinoco—D.D.
**Now attributed to Miiller—D.D.

Suborder of True Sirenians

Subordo TRICHECHIFORMES Hay,
1923

Family of Kapustniks, or Steller’s sea cow”
Familia HYDRODAMALIDAE Palmer, 1895

Genus of Kapustniks, or Steller’s sea cow

Genus Hydrodamalis Retzius, 1794

25 1794. Hydrodamalis. Retzius. Kongl. Svenska Vet.-Acad. Nya Handl.

Stockholm, 15, р. 292. Hydrodamailis stelleri.

1811. Rytina."* ПИзег. Abhandl. 4. Berlin. Akad., 1811. p. 64. Rytina

borealis.
1814. Nepus. Fischer. Zoognosia, 8, p. 640. Nepus stelleri.

1822. Stellerus. Desmarest. Mammalogie, pt. 2, р. 510. Stellerus

borealis. (У.Н.).
KAPUSTNIK, SEA COW, STELLER’S COW"

Hydrodamalis* gigas Zimmermann, 1780

1780. Manati gigas. Zimmermann. Geograph. Gesch d. Menschen und

d. vierf. Thiere 2, p. 426.
1785. Manati balaenurus. Boddaert. Elenchus anim., p. 173.

The characteristics of the family and genus are not given here, as they are clear
from the characteristics of the order and the species description given below. Only the

main generic synonyms are given.

“The first (and correct) spelling of the genus name. Rhytina is a later emendation.
''The name “kapustnik” [from cabbage] and “sea cow” are vernacular. The first
originates from “sea cabbage’”—a name which was and is still given to some seaweeds

used as animal food.
“Steller’s cow” 15 a literary name translated from a name of German origin.
*Misspelled Hidrodamalis in Russian original—Sci. Ed.

1788. Trichechus manatus, var. В. borealis. Gmelin, Syst. Nat., 1, р.
61.

1794. Hydrodamalis stelleri. Retzius, Kong]. Svenska. Vet.-Akad. Nya
Handl. Stockholm, 15, p. 202.

1815. Rytina cetacea Illiger. Abhandl. 4. Berlin. Akad., 1811, р. 70.'°
(V.H.).

Description and Biology"’

26 The body length of an adult animal exceeds 720 cm. The dimensions of
the female killed on Bering Island on June 12, 1742 were as follows'*:

“Body length from top of upper lip to end of

right lobe of caudal fin 741.5 cm
Distance from top of upper lip to mouth angle 39.5 cm
Distance from top of upper lip to shoulder 132.5 cm
Distance from top of upper lip to genital opening 490.0 cm
Length of genital opening 26.0 cm
Tail length from constrictor muscle of anal

opening to beginning of lobes of caudal fin 192.5 cm
Head circumference at level of nostrils 78.0 cm
Head circumference at level of eyes 122.0 cm
Neck circumference at occiput 204.0 cm

'6The Commander [Bering] Island is the type locality of all of these names, based
on the description of Steller.

Тре following description of the body structure and habits of the animal, written
between quotation marks and without reference to author, is a translation from Steller
(1751, 1753), done for this book.* This text is accurate and almost complete. Only a
few parts, of interest only for Steller’s contemporaries (concerning identity of his species
to the American one, the possible movement of the animal on land, its use in
medicine . . . etc.), the greater part of the anatomical section, some measurements (Steller
gave 42) and other redundancies were left out. Data of other authors are given with
reference to their sources.

The complete work of Steller, written in Latin and almost simultaneously published
in German, has not yet appeared in Russian. Only small parts of it were recast or
published in the form of free translations by S. Krasheninnikov (1755) who had at his
disposal Steller’s manuscript. Steller’s text was not always consistent. He wrote his
famous work in Kamchatka or on the return journey to Petersburg. Steller died along
the road through Siberia in Tyumen in 1746. It is possible, judging by the accuracy
with which he kept his diary (1793), that at least part of the work was written on the
island (V.H.).

'’Translated from Paris feet (V.H.).

*But see also Jordan, D.S., 1899—D.D.

Height of anterior part (proboscis) 21.0 cm
Trunk circumference at shoulder 367.0 cm
Maximum trunk circumference halfway

along posterior part of the abdomen 622.0 cm
Tail circumference at point of divergence of

lobes of caudal fin 143.0 cm
Distance between ends of lobes or width of tail

across lobes 199.0 cm
Height (thickness) of tail lobe 26.5 cm
Length of entire internal (upper) lip area which is

covered by vibrissae or lip broom 13.2 cm
Its width [lip broom] 7.7 ст

Width of external upper Пр in chin direction with

its declined surface which is entirely covered

with long white bristles 35.7 cm
Its height [external upper lip] 25.5 cm
Width of the lower lip, which is dark, without hairs
jor bristles, smooth, drooping on breast,

heart-shaped 19.7 cm
Its height [lower lip] 17.3 cm
Distance from lower lip to chest 138.0 cm
Measurement or width of mouth from one end

to other 52.0 cm
Width, or rather length, of stomach 112.0 cm
Length of intestine from mouth to anus 1521.0 cm

Accordingly it is 20.5 times longer than the
animal itself (body length is more than twenty-
four English feet as shown from the first

measurement).
Distance from genital opening to constrictor muscle

of anal opening 20.4 cm
Diameter of trachea below larynx 10.7 cm
Length of heart 56.0 cm
Width of heart 64.0 cm
Length of kidney 80.0 cm
Width of kidney 45.6 cm
Length of tongue 30.6 cm
Width of tongue 9.0 cm

Length of teats 10.26”

Di,

Fig. 2. Depiction of the kapustnik (Steller’s sea cow) Hydrodamalis gigas Zimm..,

from the chart of the first officer of У. Bering’s “Saint Peter”, Lieutenant

S. Waxell, in 1742. The drawing was probably made by G. Steller or is a
copy of his drawing (from Pekarskii, 1869).*

According to Steller the number of vertebrae is 6 cervical, 19
thoracic, and 35 others. The skeleton present in the Zoological Museum
of Moscow University has 6 cervical, 19 thoracic, and 32 lumbar, sacral
and caudal vertebrae.

The skull dimensions of adult Steller’s sea cows according to the
material of the Zoological Institute of the Academy of Sciences and the
Zoological Museum of Moscow University, are as follows: condylobasal
length (12 specimens) 638-M 673—722 mm, zygomatic width (17 speci-
mens including measurements of 8 specimens published by Stejneger,
1883—1884) 324-M 344-373 mm. The condylobasal length of the lower
jaw (14 specimens) 432-M 465—491 mm.

Males were, apparently, somewhat larger than females and accord-
ing to Stejneger (1883—1884) possessed somewhat wider zygomatic
arches.** Thus his material (8 specimens) was divided into two groups:
in one of them (probably males) the zygomatic width was larger than
half.the skull length from the upper end of the foramen magnum to the
anterior point of the premaxillae, and in the second (females), these
measurements are the same.

The length of the masticating plate (specimen in the Zoological
Institute of the Academy of Sciences) is 182.4 mm, its width anteriorly
is 80.5 mm, and posteriorly 81.5 mm. Its color is dirty, dark brown (in
living animals the masticating plate was white in color—see below)
(VF):

*Probably drawn by Plenisner; see Stejneger, 1936 — D.D.
**No specimens are of known sex — D.D.

“... The famous Clusius* considered this animal (Steller thought
that he worked with the American “manati”, V.H.) clumsy and very
ugly, because he saw only the hide stuffed with straw. The living animal
was, actually, sufficiently distinctive. However, on account of its exter-
nal appearance, and the style of movement, this animal is extremely
striking. It has so thick a skin that it resembles the bark of an old oak
tree rather than animal skin. The skin is black, rough and wrinkled as
if covered with small pebbles, or resembles a shagreen, and is hard,
elastic and devoid of hair; an axe or harpoon can hardly pass through
it. Its thickness is about an inch (25.5 mm, V.H.) and, in transverse
section and because of its color and smoothness, it greatly resembles
ebony (black, V.H.) wood. This external cover is, however, not the true
skin but rather its external layer (epidermis). On the spine the skin is
smooth or bare. In contrast to this, it is covered, from the occiput to the
tail lobes with annular wrinkles, making the skin somewhat uneven. The
sides are very rough, as if covered with pebbles, carrying many con-
vexities looking like sponges without peduncles (?). The skin looks
disgusting, especially on the head. The described upper layer of the skin
invests the whole body like a shell. Its thickness frequently reaches an
inch and consists entirely of tubules, as in Spanish rattan or in Hindu
and Chinese bamboo. These tubules adhere closely to each other as if
united, and are so perpendicularly located that they can be separated
along their whole length. Each fiber (“hair”, V.H.) lies and is fixed in
the dermis at its lower end, is somewhat rounded, swollen and thick-
ened, and has a tuberiform bulb, and therefore, the cut-off pieces of this
upper skin have [when seen] from below many swellings like Spanish
skin (Morocco ? V.H.). On the contrary, the upper surface of the dermis
lying under it is, as in a thimble, entirely covered with small depressions
in which the bulbs of the epidermis lie. Since these so-called tubules lie
very close to each other, they are always wet and swollen, but if this
upper layer of the skin is horizontally cut off, nothing is excreted from
the tubules and the cut-off surface is entirely smooth, similar to a cut
piece of claw. However, when these pieces are dried in the sun, cracks
are formed and the whole piece may be broken like a piece of bark and
the whole structure of the skin, in the form of distinctive tubules, is quite
obvious. Through these tubules, a liquid or watery mucus is excreted,
which is moré on the head and side and less on the back. If the animal
lies for some hours on the dry shore the back becomes completely dry,

*С. Clusius, 1605, Exoticorum libri decem, Leiden — D.D.

Fig. 3. Depiction of Steller’s sea cow from “Zoographia”, Р.С. Pallas (1811).
The drawing is, probably, a copy from the figure of S. Waxell (see Fig. 2) and
described by Pallas as a “draft”.

while the head and side remain wet or moistened. Thus, the thick upper
layer of the skin on the animal performs two functions: when the animal
occupies places with sharp stones, or when in winter it is always found
in the midst of the ice, it does not strip off its dermis during feeding,
or when strong waves strike the animal against rocks and submerged
stones, which I myself often saw, it does not die, but on the contrary,
is protected as with armor. On the other hand, in summer and as a
consequence of the very high evaporation, the animal does not lose its
heat and in winter, with extreme cold, it can survive. This occurs be-
cause the animal cannot stay under water for a long time, like other
water animals and fishes, but during its feeding, half its body must be
above water, and subjected to the effect of cold.

According to my observation of many dead animals that were cast
up by the sea, these animals died because their external skin layer was
damaged by rocks; this occurs mainly due to the ice in winter.

In addition to this, I have frequently noticed that when the wounded
animals were pulled by harpoons to the bank, they struggled very furi-
ously by their abdomen and tail while being supported by their forelimbs,
and thus large pieces of the external skin layer were detached. I also
noticed the detachment of these layers from the forelimbs or hands and
from the so-called hoof (hand, V.H.) as well as from the lobes of the
tail, and this confirmed my opinion .. .”

“... some sea cows had оп this upper skin layer fairly large white
spots and stripes and therefore the skin appeared speckled and this color
penetrated through to the true skin [dermis].

The upper skin layer on the head, around the eyes and ears, on the
mammary glands and arm-pits, possessed a tuberculous (stony) form or
the form of shagreen, and was thoroughly filled with harmful insects

Fig. 4. Depiction of Steller’s sea cow assembled by J.F. Brandt (1846) and

identified by him as an “imagined picture” (“Icon idealis”). The skeleton of the

animal was not known at that time and Brandt had at his disposal only fragments
of the skull and masticating plates.

which grow round the skin.’ Very frequently, the latter appeared to be
perforated by these insects, and even the true skin [dermis] was injured.
If this occurs, the oozing lymph of the penetrated glands, in which there
are usually vesicles containing fat, forms large, thick protuberances
similar to those on whales, and such patches make the body ugly.

Under this previously described upper layer of skin there lies around
the whole body the true (actual) skin [dermis]. Its thickness is two lines
(about 5 mm), soft, of white color, very compact and firm, resembling
that of a whale in respect to its structure and thickness, and perhaps
required for the same purposes.

The head, as compared to the huge body, is small, short and where
it ends is not evident; it is long and almost quadrangular in form;
however from the crown to the lower jaw, it broadens. The crown itself
is flattened, and its black superficial skin has bumps (stony, V.H.), or
looks like shagreen and appears almost broken; it is thinner (by 1/3)
than in other places, and is easily separated. From the occiput to the
nostril, the surface is bent and again bent from the nose towards the
nostril as far as the lips. The anterior proboscis (the protruding part of
the upper lip, V.H.) has a length of 8 inches (20.5 cm) and, from the
nose to the occiput, it is strongly thickened.

The shape of the mouth (opened mouth) does not pass backwards,
but is located along the sides. The external lip is so large, and declines
to the angles of the face and extends far above the lower jaw, that if
one examines only the head, he thinks that the shape of the mouth
opening extends backwards, but at a very low level.

An amphipod crustacean — Cyamus rhytinae Brandt. (V.H.).

The opened mouth is small in relation to the animal’s size; how-
ever, it need not be large because the animal feeds only on seaweeds
and lives on them.

Both upper and lower lips are paired, and divided into external and
internal regions.

The external upper lip protrudes obliquely forward as the most
anterior part of the proboscis and has a semicircular form. It is flat-
tened, swollen, thick, 14 inches (36 cm) wide and 10 inches (25.5 cm)
high, white in color, smooth and covered with a large number of tu-
bercles or cones. From the middle of each tubercle grow white transparent
bristles, 4—5 inches (10—12.5 cm) long.

The internal lip is 5 inches (14 cm) long, 1/3 inch (8 mm) thick,*
completely separated from the external one and united with the external
lip by its base only; it hangs over the palate like a calf’s tongue (!),
is all tattered and is hard, like dry twigs of a broom. The above-
described upper lip tightly closes the mouth from above, is mobile, and
is used to browse on seaweeds in much the same way as horses and
bulls collect their food by sliding the lips apart and extending them
forward.

The lower lip is also paired. Externaily it is black, smooth, without
bristles, of irregular heart-shaped form, and sometimes looks like a
chin, seven inches (18 cm) wide and six and eight-tenths inches (17.5
cm) in height. Internally, the lower lip is only slightly separated from
the external one, is rough and when the mouth is closed, it is not seen
at all because the arch-shaped external lip lies under it; however, the
internal lower lip adheres to the internal upper one and tightly closes
the mouth.

Where the lower jaw adjoins the upper one, the space remaining
between them is occupied by numerous dense and thick white bristles
one and one-half inches (38 mm) long. This prevents anything from
being dropped from the mouth during mastication, or washed off by sea
water: the water, which together with food is always taken into the
animal’s mouth, is drained outside when the mouth is closed.

The above-described bristles have the thickness of a goose feather
quill, white, hollow, and with follicular swellings from below. By them,
it is clear to see without a magnifying glass how nature forms our hairs.

When the animal lies directly on its abdomen, the height of the
anterior part of the proboscis or muzzle, measured perpendicularly

*From Steller (1753)? Steller (1751) reads “25 inches wide’—D.D.

from the nostrils to the lips, is equal to eight inches (20 cm); it extends
from the nose to both the anterior-most part of the lips, and to the
lateral side of the upper jaw, like mesenteries (in folds), anteriorly
rounded, then thickens and its size greatly increases. The external lips
are very greatly enlarged, thick, as Ш swollen and, as in cats, carry
many white pores (!) or orifices. From each, thick white bristles protrude
which become thicker as they gradually approach the mouth opening.

The thickest bristles are those growing between the lips of both
jaws. They use these bristles in browsing seaweeds, and they prevent
anything from falling from the mouth. The lower jaw is shorter than the
upper one and is movable. The lips of both upper and lower jaws are
mobile and function as in our draft animal [horse]. When the animal
picks vegetation with its hands or forefeet from the plants growing
underwater on the stony bottom, it cleans the fronds from the hard stems
and roots (which are not utilized as food) with the above-described
bristles as if cut off with a blunt knife. These stems and roots are thrown
up by the waves during [high] tide and heaped on the shore in very large
piles, indicating that an animal is present. Since the stems of aquatic
plants are much harder and thicker than those of terrestrial plants, these
beasts must have harder, more compact lips than any terrestrial animal.
Therefore, such lips are not utilized by man as food and they are not
softened by boiling or any other means. The inner lips are so con-
structed that, if they are cut open, separate small areas are seen on them
similar to those on a checkers or chess board. These are innumerable,
very small and thick red muscles of rhomboidal or trapezoidal shape.
Between these muscles numerous areas resembling tendons are located, —
and form a net or tissue of vesicles filled with liquid fat. These lips
easily release all their fat when cooked in boiling water and when they
are defattened, all the white areas look like the numerous tendons of a
net. This natural structure, it seems to me, has three purposes, namely:

1) in order to make the lips stronger and more compact and protect
them from external injury;

2) ш these muscles the heads and tails, i.e. their beginnings and
ends [origin and insertion] are so located that their heads are slopingly
directed toward the mouth opening while their tails, or ends, are located
obliquely in relation to the parietal, in such a way that their beginnings
and ends form a sort of garland and this helps raising and moving the
heavy lips;

3) with the assistance of this structure, the lips can be, to a certain
degree, bent in a helix or spiral and since the whole head is covered by

a thick cortex it bends with difficulty and thus, when the animal wishes
to browse on elastic seaweed, it is not necessary to move the whole
body.

Mastication in these animals occurs in a manner different from
other animals. It is achieved not by teeth, because they possess no teeth,
but with the assistance of 2 long bones found in their place. These bones
[masticating plates] are large, of white color and are considered as
whole rows of teeth or as one tooth mass. One of these bones is fixed
to the palate, and the other to the lower jaw.

These bones are, however, attached in an entirely unusual manner
and the method of their attachment cannot be attributed to any of the
known ones. They cannot be described as fixed because they do not
insert in the jaw as do teeth, but their protuberances and pits lie in
corresponding protuberances and pits in the palate and lower jaw.

Fig. 5. Depiction of Steller’s sea cow made from the so-called “Royal Agriculture
Manuscript” of S. Waxell and his later (compared to that described by Pekarskii,
1869—-see Fig. 2) chart (Bikhner [Biichner], 1891). In the original, the upper
drawing has a length of 26.5 cm (in that of Bikhner, 1891—from the nose to the
engraved tail 20 cm), and two times more than the lower drawing. The upper
drawing (“Royal Agriculture Manuscript No. 2”) is, apparently, a more detailed
drawing of the same animal in Fig. 2. The lower drawing (“Royal Agriculture
Manuscript No. 1”) was greatly changed during re-drawing, probably by the same
source (according to Bikhner, 1891—4115 is a depiction of another animal).

Moreover, the dental bones are fixed in the anterior part of the hard skin
of the inner upper lip, along the sides of the mouth in the edge of the
bones, and are finally inserted into the palate and lower jaw in the
posterior part by means of a double outgrowth, and that is how they are
attached.

These grinding bones occupy the place of all molar teeth and, like
a thimble or sponge, have many small holes on the lower surface through
which, as in the teeth of [other] animals, the arteries and small nerves
pass dorsally; these dental bones are far from smooth and covered with
numerous bent wavy grooves; between them prominences are raised
which, in their turn, occlude during mastication, into the grooves of the
opposite dental bone and between them, and the marine plants are
ground as in a hand or fulling mill.

The nose lies on the uppermost part of the head and to the sides as
in a horse; the two nostrils are separated by the nasal septum of one and
one-half inches (38 mm) in width. The nostrils themselves are 2 inches
(51 mm) long and the same in diameter; they are open, but inside, there
are many flexures or blind pockets. The inner walls of the nasal open-
ings are extraordinarily thick, wrinkled and covered with compact skin
with many black pores which have bristles protruding from each of
them. These bristles have the thickness of sewing thread and are one-
half inch (13 mm) long and are easily detached; they perform the function
of those in other animals.

The eyes lie exactly mid-way between the end of the muzzle and
the ears, at the same height as the upper part of the nose or slightly
higher. They are relatively small in comparison to the large body of the
animal, and are not larger than those of sheep. Externally, they have no
eye-lashes or any similar structure surrounding the eyes, and they pro-
trude from the skin through a rounded hole, which according to my
measurements, has a diameter of scarcely one-half inch (13 mm). The
iris is black in color and the eyeball is yellow blue. The corners of the
eyes are not noticeable; moreover, when the skin around the eyes is cut
off, a cartilaginous body like a cock’s comb protrudes at the large
(anterior) corner, as in the case of the sea otter. When needed, this body
closes the eye, nearly in the same way as in animals which are grazing,
i.e. searching in dusty and dirty soil, and has such a membrane (third
eyelid, V.H.), which can be stretched by the animal on the eyes to
prevent the entrance of any harmful objects. This cartilaginous body
forms, with its other side, the wall of the lachrimal sac and is connected
with it by a common compact skin. On cutting into the lachrimal sac,

AOYPPUOY ‘Мм Aq Sunureg ‘ии 572818 sppUMpO.PAL] ‘MOD BIS $.19191$ “| Эа

dense mucus is found inside; the lachrimal sac is large enough to allow
the entrance of a chestnut, and it is lined by glandular skin.

Externally, the ears are represented by a small, open orifice but
there are no traces of external ears. The ears themselves are unnotice-
able, unless a person searches carefully for them. They cannot be
differentiated from the surrounding skin which resembles shagreen. The
shaft of a feather can hardly pass into the ear.

The internal ear canal is smooth and coated with an extraordinarily
smooth (polished) black skin, and therefore it can be easily detected,
when the muscles of the posterior part of the head are separated, by its
color.

The tongue is 12 inches in length (30.5 cm), its width three and a
half inches (9 cm), and as in cows, it is pointed at its end; its surface
is rough as a file and covered with low warts. The tongue is so deeply
placed in the mouth that one may doubt its presence. If it is pulled by
hand from the mouth, its anterior part fails to reach the grinding plate
of the mouth by one and one-half inch (38 mm). However, if the tongue
were as long as in other animals, it would interfere with the above-
mentioned wide bones during mastication.

The head is not visibly demarcated from the neck, and these are
similarly connected to the trunk without any demarcation in any place,
and thus the body is constructed as in a fish. The neck is not clearly
demarcated, except that it is half the length of the head, elongated,
round, and somewhat more slender than the posterior part of the head.
In spite of that, the neck is movable due to the presence of movable
vertebrae. However, these movements are only observed during feeding
in living animals. During this, the animal lowers its head as does an ox,
grazing in a dry pasture. However, if the animal lies resting or after
being killed, the animal seems to have an immovable neck owing to its
thick clumsy outer skin and especially due to the absence of any exter-
nal trace of the cervical vertebrae.

From the scapula to the umbilicus, the trunk is again strongly thick-
ened; and from the latter to the anus, the trunk size decreases and it
again becomes slender. The flanks are as bulging as the belly itself, and
the latter, due to the great weight of the intestine included in it, is very
swollen, stretched and distended by the intestine. From the umbilicus to
the anus, the trunk is thinned to the same degree as from the chest
(mammary glands) to the neck.

In spring and autumn, when the sea cow is in fat condition, the back
is somewhat convex, and in winter when they are emaciated, the back

appears flat, and on both sides of the vertebral column there appear deep
hollows; and thus the vertebral column and vertebrae are clearly seen.

The ribs are curved up on both sides of the spine as a convex arch
and in those places where they articulate with the vertebrae, as in the
case of human beings, they again bend down forming an arch, where
on both sides of the vertebral column they form a double groove along
the whole length of the back.

The tail starts at the 26th vertebra; the caudal vertebrae are 35.
Posteriorly from the anus and in the direction of the tail fins the tail
becomes thinner and not completely flat, but rather somewhat quadran-
gular . . . the tail muscles fill the empty places in the vertebrae and the
angles between their processes by 4 plates and therefore, the tail looks
like a long quadrangle with blunt angles.

The tail is generally thick, extremely strong, very hard, stiff (slightly
movable) and ends in a black fluke. The latter is not divided into rays
but forms a single unit. Its material is similar to the whalebone (baleen)
which is used by the dressmakers, and, similarly, is composed of leaves
lying one above the other as rings of a tree or the grain of a piece of
wood. At a distance of one-quarter from the end, it appears wider and
somewhat resembles a coarse spindle*.... The fluke itself, forming
the tail end, has a width or length of 78 inches (199 cm), a height of
7 3/10 inches (18.5 cm), and a thickness of 1.5 inches (38 mm)... .

The caudal fluke somewhat resembles pincers (?), with horns or
hands of equal size. . . . The tail blades are located, as in whales and
dolphins, to the sides, and are not, as in the majority of fishes, in a line
with the spine. If the sea cow slowly moves its tail to one side, it slowly
swims forward, but if it strongly moves the tail upward and downward,
it rapidly swims forward and tries in this way to move away from
harmful hands wanting to pull it on to the shore.

The most remarkable thing in which this animal differs from all
terrestrial animals which live in water.and from aquatic animals which
spend some time on land, and from all animals in general, is its entirely
unique hand or fore limb, if it can be so called. At the neck, two limbs
originate straight from the shoulder bones, 26 inches (68 cm) in length
and consisting of two parts. The uppermost bone [humerus] is con-
nected to the scapula by a flat articulation.

Here, as in the case of humans, there are radius and ulna, both of
which border on the carpal bones with the metacarpal bones (called

*In Russian original, “‘osti kolosa”; literally, grass awn—Sci. Ed.

metatarsals and premetatarsals, V.H.), but neither fingers, nails nor
claws can be detected. The above-mentioned anterior and middle (meta-
tarsal and premetatarsal) parts of the limb are covered with dense fat
and coated with many ligaments and superficial skin, and have the form
of the newly overgrowing skin on a wound on the human extremity.
However, the skin and especially its superficial layer is thicker on that
part and also harder and drier, so that the ends of the hands roughly
resemble a kind of crustacean claw [nail] or horse-hoof, even though
the horse-hoof is sharper and narrower and therefore more adapted to
digging, or scraping. At the posterior border, these limb ends are smooth
and curved; from below, they are, to a certain degree, concave (empty)
and covered with numerous dense bristles about one-half inch (12—13
mm) in length and, when scratching, are as sharp as the twigs of a
broom. In one of these animals I noticed nipper-like limbs, divided into
two parts, similar to the cow hoof. However, such division was not an
actual one and was restricted to the superficial skin layer, and was
probably accidentally brought about and not naturally created in this
form. This is still more possible, as the superficial skin layer lying on
the so-called claw handle is very dry and may be easily cracked.

This Platonic man, as the famous Ratyus (Ray, V.H.)* said in jest,
used its hands in different manners. It can use them as fins in swim-
ming. It walks on them in shallow waters as on legs; it holds itself upon
them, resting among the slippery submerged stones and rocks, and
scrapes and tears off the sea plants and seaweeds from the rocky
bottom as do horses with their forelimbs. It can lean upon them and
thus make great resistance when wounded by a harpoon and dragged
from water to the shore. In such conditions, it holds so strongly that the
upper layer of skin on the hand is split and torn off in pieces. During
sexual intercourse, the female lies on her back and swims in this
position. The male lies upon her; she holds the male strongly and he,
in his turn, embraces her.

On the anterior part of the breast, two teats are found, located in
a manner different from that in other animals. Each one lies under the
corresponding arm as in the case of humans and has a similar appear-
ance. Each breast has a length of 1 1/2 feet (45 cm), is convex, and
bears many folds or spiral flexures directed inwards. The breast is
glandular, extremely hard, harder than the cow udder and without fat
deposits. The fat layer covering the whole body is found also on the

*John Ray—D.D.

Fig. 6. Skull of Steller’s sea cow. Upper figure, side view of skull; lower,
dorsal view (from Stejneger, 1884).*

breast and is as thick there as on other parts. However, the upper layer
of skin is thinner, softer and more wrinkled. The nipple also is of dark
outer skin, with circular wrinkles. However, this superficial skin layer
is soft; the nipple hangs under each hand or shoulder; in lactating
animals, it has a length of 4 inches (10 cm) and a thickness of 1.5
inches (38 mm). If the lactation period is over or if the animal has not
yet given birth, the nipples do not protrude and are so short that they
can be considered as accidentally occurring warts. At that time, the
breast also is not very swollen. The milk of these sea cows is as sweet
and fatty as regards its taste and density, as that of sheep. Frequently,
I drew milk from nipples after the death of the animals, as from cows.
The area around the nipple is very wrinkled and a little raised above
the surface of the breast. In each nipple there are from 10 to 12 milk
ducts which open into the teat. The boiled mammary glands were
somewhat harder than a cow’s udder and they had a weak gamy taste.

*Actually from Simpson (1932) — D.D.

These animals mate in the same way as human beings. The male
lies above and the female lies below. The penis 15 32 inches (81.5 cm)
in length; it is tightly fixed to the belly wall by its sheath, reaching the
umbilicus; it is thick and ugly in its external appearance. It is similar
to that of the horse and has the same head, but is somewhat larger.

The female genital opening is located eight inches (20 cm) in front
of the anus. Anteriorly, the opening is nearly triangular . . . to the rear
it narrows. ... The anus... 1s closed by a constrictor muscle, not very
tightly. Its width is four inches (10 cm); the sphincter is white... .”

From the biological observations of Steller given in the part of his
work entitled “Description of behavior and nature of this animal”, the
following lines deserve particular attention (many interesting observa-
tions are not directly related to the biology of this beast). “. . . I was
lucky enough, or rather unfortunate, to be obliged to observe the be-
havior and customs of this animal in front of my door for ten months”
and thus I wish briefly to state that I can explain [their behavior] with
trustworthiness.

These animals like shallow and sandy places on the shore. How-
ever, they very willingly stay where rivers and creeks pour into the sea,
and on their banks, as they are attracted to the fresh water coming to
the sea; therefore, they stay there in groups. When they feed, they lead
their young or the newly born ones in front of them, following them
from behind and on the sides, and very carefully surround them, keep-
ing them in the center of the herd. When the tide rises, they come very
near to shore, so that I could not only beat and prick them, but some-
times even stroke them with my hand on their back. If they suffer from
a severe pain, they only moved away further from the coast than usual,
but they soon forget that and come near агат.”

Usually, their whole families are found close to each other; the
male with the female (“with his wife”), with one adult [offspring?] and
also some other young. Each husband, apparently, does not have more
than one wife; they give birth to young at any time of the year, but most
often, judging by the newly born young, in autumn. Proceeding from

**Bering’s ship ‘Saint Peter” escaped a rock and was very lucky to be thrown on
the flat shore of a broad bay, where the crew spent the winter in mud huts and where
Bering himself died. This place was very favorable habitat for sea cows (V.H.).

*|Steller writes elsewhere “‘. . . it is rather possible to believe that it 15 possibie to
make this animal tame, rather than this strange cunning which is attributed to it in
stories (spoken about the American species, which Steller considered the kapustnik,
V.H.). Due to its extraordinary stupidity and greediness for food, this animal is tame by
nature, and there is no necessity to domesticate it. . .”.

Fig. 7. Bering Island, shore near Cape Manati. Former habitat of Steller’s sea cow.
Photograph by S.V. Marakov.

the fact that I observed their mating early in spring, I think that preg-
nancy lasts more than one year, and that they give birth to no more than
one young, which can be deduced from the short uterus and the fact that
the adult female has only one pair of nipples. Moreover, I have never
observed more than one calf beside its mother.

These insatiable animals eat continuously. They are extremely greedy
for food, so they hold their heads all the time under water; accordingly,
they very rarely worry about their life and safety, and it is easy in a
boat or even swimming naked among them to choose which one 1$
needed to remove from the sea. During feeding they do nothing but
thrust their nose out from water every four or five minutes, and, making
a noise similar to the neighing, sniffing, and snorting of horses, they
breathe out air with a small amount of water. During feeding, they put
one leg after other and in this way they swim slowly, or walk like oxen
or sheep. Half of the trunk, mainly the back and flanks, are always
protruding from the water. During feeding, sea-gulls sit on their backs
and pick the lice (crustaceans, V.H.) found on the upper layer of the
skin and in this way they are useful to them, in the same way as crows
do on swine and sheep. The sea cows do not eat all the types of sea
weeds without selection, but they mainly eat: 1) wrinkled cabbage, like
Savoy cabbage with latticed leaves; 2) water plants with upper parts
similar to a cudgel; 3) plants which look like the whips of ancient

Romans; 4) very long sea weeds with a wavy vein edge.” In places
where these animals were feeding, even for one day, the roots and
stems thrown out by the sea lie on the shore in large heaps. After they
feed to satiety” some of them lie on their back. To avoid being on dry
land during ebb-tide, they swim away from shore farther into the sea.
In winter, these animals frequently die under the ice which floats along
the coast, and dead animals are cast up onto the shore by the sea. The
same also happens when the waves strongly strike the rocks, taking
these animals with them. In winter the animals are so emaciated that
all their ribs can be counted along the vertebrae.

In spring, they come together as do human beings, mainly in the
evening when the sea is calm. However, before copulation they show
many amatory signs in their relation to one another. The female slowly
swims in the sea here and there, and the male follows her all the time;
the female moves around for a long time near the male until she reaches
extreme sexual desire. She then lies on her back as if she is tired and
does that seemingly unwillingly. When this occurs, the male almost in
a frenzy rushes on her, and they copulate with each of them embracing
the other.

Capture is carried out with the help of a large iron hook, one end
of which looks like a ship’s anchor (having a hook directed back-
wards); while the other end has an iron ring to which a very long and
strong rope is connected. A very strong man held this hook (harpoon
V.H.) and sat with four—five persons aboard a boat. One of them
handled the tiller while the other three or four rowed; and with them
he carefully approached a whole herd of these animals. The hunter

According to the opinion of A.V. Vozzhinskaya (Institute of Oceanology of the
Academy of Sciences), Steller wrote about the following species in order: Agarum sp.;
Nereocystis sp.; Alaria (fistulosa); Zostera, or Laminaria longipes.

*3Steller writes elsewhere that the intestine of these animals was always completely
filled with large amounts of food: “Те stomach is astonishingly large, its length is 6
feet (180 cm) and width 5 feet (150 cm); it is filled with food and seaweeds to such
a degree that four strong men with great difficulty can scarcely drag it from its place
with a rope.” ,

The animal’s feces “Бу its form, amount, odor, color as well as all other essential
characteristics is recognized by the experienced stable-man as that of a horse’’. Steller,
in his first days on the Island, considered these feces which were thrown up by the sea
as horse feces, and considered this as a proof of his view, that America was not far from
this place (no horses were present at that time in Kamchatka) (V.H.).

“... One of these animals was stranded on dry land because it slept and the sea
at that time receded, so it was not able either to go back or to resist. It was killed with
cudgels and axes.

3637 ~— Fig. 8. Bering Island. The coast on which Steller’s sea cow was found.
Photographs by S.V. Marakov.

stood on the anterior part of the boat holding the hook in hand; and
when he was able to hit the animal, he did. Thirty persons who were
standing on the shore holding the end of the cord tried to hold the
animal and with great effort pulled it to the shore; the animal, at that
time, resisted with fear. Those who were on the boat, held their place
with the help of another rope, and continuously beat and stabbed with
knives and other sharp instruments to such a degree that the weakened
animal, completely exhausted and worn out by the wounds, was drawn
up on the shore.

Some [people] cut large pieces of meat from the animal while it was
still living. During this, the animal only struck violently with its tail
here and there, and so strongly resisted with its hands that large pieces
of the superficial skin layer were usually detached from them. More-
over, the animal was deeply breathing and moaning. Blood shot upwards
as a fountain from the wounded back; so long as the head of the animal
was hidden in water, there was no bleeding, and when it took a breath
in air (for this reason it raised its head from water) bleeding began

Fig. 9. Peregrebnaya Bay near Cape Manati on Bering Island—the former habitat
of Steller’s sea cow. Photograph by S.V. Marakov.

again. This is because when the back was wounded, the lungs which
lie under it were also wounded and the more frequently the lungs were
refilled with air, the more frequently they expelled blood....

The adult and very large animals are more easily hunted than
calves, because the latter swim more rapidly and vigorously. If a har-
poon was thrust in them [large ones] and was firmly attached to them,
it remained in their bodies and was held there, but the calves sloughed
off their skin and escaped, as was observed many times.

If the harpooned and captured animal begins to struggle strongly,
only the animals in the herd swimming nearby come close to it and try
to help it. Thereupon some animals try to turn the boat upside down
with their backs, while others lie on the rope” and try to cut it off or
to strike the rope with their tails to pull the harpoon out of the back of
the injured animal, and this they sometimes succeeded in achieving.
Definite proof of their abilities and at the same time, of their conjugal
love was shown in that one male whose female was captured
by a harpoon and pulled out onto the sea shore, tried with all his strength
to help her, but his attempts were unsuccessful, and in spite of all the
blows which we dealt him, he followed the female to the shore, and
several times suddenly threw himself like an arrow toward her although
she was already dead. The next day, as we were getting ready early in
the morning to cut the dead animal into pieces, we found the male still
standing beside his female. Even on the third day, when I came to
investigate the internal organs in detail, the male was still there.

Concerning its voice, this animal is dumb and does not produce any
sound. It only breathes loudly and also moans when wounded. But
concerning sight and hearing, nothing can be said. These animals al-
ways hold their heads under water and therefore they cannot hear or see
well. This gives the impression that the animal itself has no need for
these senses.

. . . Several times it occurred that dead animals were cast up by
storms around Cape Kronotskii, which is called the Nose, and Avacha
Bay [on Kamchatka]. Because of the type of food which is consumed
by these animals, the inhabitants of Kamchatka called these animals
“Kapustnik” [cabbage-worm] in their language, which means eaters of
weeds; I learned of this, when I returned (from Bering Island, V.H.) in
1742.

*5According to the description of Bering’s companion, Sofron Khitrov, rope of 4.5
inch diameter, i.e. about 100—110 mm was employed (Pekarskii, 1869).

. . . The fat layer under the upper skin layer and the dermis
covering the whole body is as thick as a palm’s width, and in some
places, three-quarters of its width. The fat is full of glands (!), soft,
white in color, but in sunlight it turns yellow, like May oil. The odor
and taste of the fat is pleasant and cannot be compared with that of any
other marine animals, and it even exceeds that of tetrapod animals.
Moreover, this fat can be kept for a long time even on very hot days,
for it neither spoils nor smells. After melting it 1s delicious and tasty,
and is preferred to any other oil. Its taste is almost like sweet almond
oil and it can be used for any purpose for which any oil is used. It burns
brightly in lamps and produces no smoke or bad smell. . . The fat of
the tail is harder, stiffer and therefore it is more delicious when cooked.
The meat has stiffer and thicker fibers than beef; it is also redder in
color than meat of terrestrial animals and it is surprising that the meat
of this animal can be kept in the very hot days for a long time without
producing any smell, even if it is everywhere full of maggots[. . .]°°

The meat of this animal, it is true, requires a longer time for
cooking but when cooked, it has.a superior taste and cannot be differ-
entiated from beef. The fat of young sea cows is similar to that of fresh
swine lard and it is hardly differentiated from the latter. Moreover,
their meat cannot be differentiated at all from veal, and it also is
cooked rapidly; during cooking, it also swells like young pork meat and
occupies in the pot twice the space as originally.

Fat in deposits, as in the head and tail, is viscid and therefore can
scarcely be boiled. The most preferred muscles are those of the lower
surface of the abdomen, back and flanks. Such tallow is not only fit for
salting—which many persons doubted—but when salted it 1s like that
of corned beef and has a very nice taste. Internal organs—heart, liver
and kidneys—are very hard and we did not utilize them because we had
enough meat, for a mature animal weighs nearly eight thousand pounds
which equals éighty centners, or 200 Russian poods (3200 kg). The
number of these animals was large on this single island, so the inhab-
itants of Kamchatka might continuously feed only on them.

A particular harmful insect*, like a louse, causes much trouble to
sea cows. They stick in large masses on the wrinkled limbs, on mam-
mary glands, teats, in hidden places, anus and in the shagreen-like

6The explanation given by Steller of this phenomenon is of по interest, but the fact
itself of the possible long-distance transport of bodies by the sea, especially to Kamchatka,
is significant (V.H.).

*See footnote 19, р. 24.—Sci. Ed.

cavities of the upper skin layer. . . . These insects pierce both the upper |
layer of the skin and the dermis and protuberances are formed from the
released fluid, and here and there visible warts. However, these insects
are pursued by the sea-gulls which sit on the back of the animal and
pick them up daintily. Thus they offer a pleasant service to the animals
which suffer from these harmful insects .. .”.

Geographic Distribution and History of Extinction

Steller discovered the kapustnik in 1741 on Bering Island. Later, it was
established that this animal lived also on Mednyi [Copper] Island, but
information about this is very scanty. In the historical period, sea cows
were not found anywhere else. The sea cow is completely analogous to
Steller’s [Pallas’s] cormorant (Phalacrocorax perspicillatus Pall.),
which was also limited, apparently, in its distribution to the Com-
mander Islands and also was exterminated somewhat later—in the middle
of the nineteenth century.

It seemed that the sea cow must have been living in the Aleutian
Islands, where the living conditions are actually the same as in the
Commander Islands. The standing crop of sea weed is also great there,
and the same species grow as in the Commander Islands; generally,
they comprise one geographical and zoogeographical region. This par-
ticularly applies to Attu Island, the most westerly of the Aleutians
(Blizhniye group) and the nearest one to the Commander group (about
350 km Нот Mednyi Island).

Moreover, Steller did not refer to this species in places east of
Bering Island in any of his works, including the diaries, that were
published by Pallas (1793) after his death. There is also no record in
the descriptions of any other voyages in the eighteenth century. As a
result of a special collection of information and complete analysis of
the question of the geographic distribution of this species, [von] Baer
(1838)* noted the complete absence of this animal at any time in
Russian America, including the Aleutian Islands, especially the Lis’ie
and Andreyanov islands.

Later, Stejneger (1883), in his evaluation of all information (tes-
timony of Aleuts from Attu, data on Semitki and Agattu islands, as well
as observations of Turner, Voznesenskii and others), denied the argu-
ments, which are actually very vague, concerning the existence of
Steller’s sea cow on the western Aleutian Islands including Attu.

*Actually published in 1840—D.D.

He denies the existence of sea cows on Attu and [the report of]
Nordenskiéld (1882), based as it was on unwarranted dependence оп
testimony of inhabitants. A single rib found on Attu by Voznesenskii,
if it actually belongs to this species, came from the body of a dead
animal cast up by waves,”’ as Brandt (1846) correctly concluded.

In American literature, including the latest (Murie, 1959; R. Rausch,
1960), there is no evidence of the occurrence of Steller’s sea cow
around Alaska or on the Aleutian Islands, or any discovery of its
remains (not taking into account the case given by Voznesenskii).
There are neither archeological nor paleontological remains of this
species in the American territories (information from Ch.A. Repenning,
paleontologist, U.S. Geological Survey; only known are the remains of
a closely related genus Halianassa, which is not younger than middle
Pliocene, from the eastern part of the Pacific Ocean). In fact, if we
leave aside possible historical causes, it is very difficult to imagine the
emigration of a herbivorous animal which is so closely associated with
the seashore and which has very little vagility, across 350 km of open
ocean from Mednyi to Attu islands.

Sea cows were never seen in Kamchatka and were generally absent
on the Asiatic shores. Neither Steller (1751, 1753, 1781, 1793) nor
Krasheninnikov (1755) made any reference to such an occurrence. All
of Krasheninnikov’s information (1755) was related to Commander
[Bering] Island-and, as mentioned above, was based оп Steller’s manu-
scripts, which were at his disposal. There is direct information about
the absence of this species in Kamchatka in the eighteenth century. The
situation wherein some inhabitants of Kamchatka knew about this
animal which they called kapustnik (“cabbage-worm’’) 15 explained by
the washing up of bodies (see footnote on p. 40) and finding in their
stomach after dissection of a large mass of “sea cabbage”, as was cor-
rectly noted by Stejneger (1883) (see footnote on p. 35; field cabbage
was not known at that time in Kamchatka). The distance from the
nearest point on Kamchatka (Cape Kamchatka) to Bering Island is 150
km, a distance across which a carcass could be transported but cer-
tainly insurmountable for a living animal. The absence of sea cows
along the Asian coast was emphasized by [von] Baer (1838), who
showed especially that this animal was absent not only in Kamchatka
but also in the Kuril Islands.

Later, Ditmar (1900) referred to the absence of sea cows on
Kamchatka in his 1852 material. Finally, it is known that in the middle

’This rib could have been brought by workers who were preparing meat in the
Commander Isiands (V.H.).

Fig. 10. Tundra Bay on Bering Island—a place where sea cows occurred.
Photograph by S.V. Marakov.

of the eighteenth century, commercial expeditions to the Aleutian Is-
lands and Alaska, for trade in beavers, [sea otters] and other fur-bearers,
stopped off especially at Commander [Bering] Island to prepare sea
cow meat and even wintered on the islands for this purpose. This was
done because this animal was not found on Kamchatka (Pekarskil,
1869; Ditmar, 1900 and others; see below).

Some reports of the presence of sea cows on the Asiatic coast are
found in the literature, but they are false. Thus, the reports of the
observation (1762) of sea cows at Cape Shelagskii and between it and
the Kolyma, in the ice of the East Siberian Sea (Grekov, 1958, from
a document of Shalaurov), were based on some strange misunderstand-
ing. Such a casual remark as Nordenskidld’s (1882)* in which he says
that “one Russian”, around 1910, saw a dead sea cow supposedly
washed up by the sea at Cape Chaplin (north of 64° N. Lat.) is not
trustworthy. Finally, in the light of what has been said above, and as
all the data on the biology of the sea cow make absolutely clear,
“observations” of this species at Cape Navarin in the Bering Sea (about

*Actually attributable to Sverdrup (1930}—D.D.

62° М. Lat.) in 1962 (Berzin, Tikhomirov, Troinin, 1963) have no
value at all (Heptner, 1965, see below).

The history of the extinction of the sea cows is considered the most
tragic history of the destruction of a most remarkable animal in a very
short period, and at a time when the technology of animal destruction
was at a low level. Steller’s sea cows were in the true sense of the word
eaten up by human beings.

As is clear from Steller’s description and from other information
(see below), the meat and fat of sea cows were characterized by their
excellent quality. This fact was sufficiently widely advertised by Steller,
who returned to Kamchatka in autumn 1742. Moreover, he suggested—
this probably was a mistake—that the number of sea cows around the
island was very large. In fact, these animals were, apparently, numer-
ous only where the crew of the “Saint Peter” was shipwrecked and
spent the winter, and in some other especially favorable parts of the
coast. The total number of these animals could hardly have exceeded
some hundreds.

What is then the cause for such intensive destruction of this animal,
which lived around the shores of two similar, small islands lying far out
in the ocean, and which had been just discovered and were uninhab-
ited? On the one hand, Steller wrote diligently about the quality of the
meat and fat of the sea cow, and especially about the fact that Kamchatka
at that time could not supply with foodstuffs the numerous commercial
ships which were sailing from Petropavlovsk and Bol’sheretsk in
Kamchatka to the east and northeast. The Commander Islands therefore
became a base for obtaining provisions for the crews of these ships.
The system of hunting was constructed surprisingly rapidly and its
organization was very flexible, and the “profusion” of the island was
rapidly appropriated. In the next year after Steller’s return, 1.e. in 1743,
the ships which were sailing to trade in the Aleutian Islands and in
Russian America went at first to Bering (“Komandor”) Island and
worked there preparing sea cow meat, overwintering because of this.
Later this was continued by all the numerous commercial ships directed
from Kamchatka to the east for hunting arctic foxes, fur seals and,
above all, Kamchatka beavers [sea otters].

The Marine Department in Petersburg in 1765, apparently in ques-
tioning the merchant Vasilii Shilov about the distant lands situated
beyond the Commander Islands, and about the trade and geographical
discoveries conducted there by the commercial companies, asked him
particularly about the “means” that they had in those far and long

voyages “. . . how you were saved from hunger, because the food
prepared in Kamchatka was for two or three months, and flour was
especially insufficient?” The answer declared that the “trading compa-
nies at present are far from Kamchatka beyond Commander and Mednyi
and near to the North, and of others, several islands became better
known far to the East”, however, Kamchatka “cannot provide such
distant routes with provisions, especially for prolonged residence there
for hunting, and thus they used, except for flour, the abundance of
Commander and Mednyi islands”. From Kamchatka, the ships took
besides flour, only such provisions and fresh water as helped them,
even in unfavorable weather, to reach the Commander Islands “. . . and
there they spend the winter; and during the whole wintertime they are
engaged in hunting, and on the distant routes they prepare provisions
which mainly consisted of sea cows, whose meat is not only wholesome
and in great quantities, but can also be used instead of bread when
dried. In this animal there is much fat to be stocked in small barrels and
used in meals instead of oil, and one can drink it harmlessly like the
best [olive] oil. It can also be used for making fire instead of candles.
The skin of these sea cows is used instead of boards in the planking
of boats...” (Pekarskii, 1869).

Hunting itself was, on the whole, performed in the same way as
that of the people on Bering Island, and it was based on the same
biological characters of the animals, 1.e., on their habit of coming to the
shallowest water at ebb tide. However, some changes appeared in the
technique. Peter Yakovlev describes it in the Nizovtsovaya harbor,
where “. . . in the sea, near the shore, these sea cows were seen singly
or in groups. In hunting they go in a boat, eight in a group, of which
one hunter stays near the stern in front of the helmsman holding a long
pole armed with a wide, long sword-like band (called “pokolyuga”),
and the rest of the hunters sit as oarsmen. When they find themselves
in front of a sea cow’s head, they turn the boat with the stern towards
the sea cow and the hunter with the “pokolyuga” inflicts a severe
wound in that beast’s heart. After that the rowers must row quickly
away from the sea cow in order to get out of that wounded sea cow’s
reach, for it can ruin their boat by its tail or fluke . . . with such a
wound in its heart, the cow does not swim in the sea for a long time.
It soon gets tired, stops and turns with its belly up. The hunters pull that
sea cow by a hook [harpoon, V.H.] towards their boat, tie it to the stern
of the boat and row to the shore. The meat of one sea cow was enough

for 33 men for a month. The net weight of one sea cow’s meat and fat,
without bones, reached 200 poods” (Pekarskii, 1867).

Moreover, as Vasilii Shilov describes, workers used the skins of
sea cows in covering the frameworks of boats instead of covering them
with wooden boards. From Kamchatka, they took “the ready-made
wooden boat keels and the stern and bow installations for one or two
boats” and covered them with sea cows’ skins in the Commander
Islands, “because” he notes, “such large skins are not present in
Kamchatka” (this also proves the absence of the above-mentioned species
on the continent). Sewn skins (that is, from animals of smaller size) are
not convenient because they leak.

Shilov further stated that “though the ship-chandlers take with
them all the prefabricated parts, until they are actually needed (almost,
V.H.) they do not cover them with the skins but prepare excess skins
(my emphasis, V.H.). Nevertheless, they store up skins in a great
quantity and when necessary they gathered the boats in a short time,
and covered them with skins after soaking them in water... .” These
skins were, apparently, of a great value because the boats were safer
in the breakers, and what is more important, they were comfortable in
handling and very light in comparison to wood-planked ones. “A boat
covered with leather carried the same number of people (as a wood-
planked one—20 men, V.H.) and it could be pulled out onto the shore
or from the breakers, like a bladder, without risk, and could be carried
a good distance by only four men... .”

As shown, Steller’s sea cow was used by pioneers in developing
Kamchatka and Russian America rather fully, but too excessively. As
a result, it was quickly exterminated and in 1768, that is only 27 years
after its discovery, the last one was killed on Bering Island. The ani-
mals around Mednyi Island, which were, undoubtedly, fewer, were
exterminated earlier; at any rate, in 1754 there was not a single one of
them.” It is possible that on Mednyi Island one or two outlived the
indicated date for a year or two; however, in the seventieth decade of
the eighteenth century nobody saw them and Bragin, who was spending
the winter on Bering Island in 1773—1774, makes no mention of sea
cows. There are no later records of them (Berg, 1946).

8Steller (see above) apparently considered that 200 poods is the total weight of the
beast (V.H.).

**Peter Yakovlev, sent by the government according to the suggestion of M.V.
Lomonosov, to Mednyi Island to search for useful minerals (copper), whose whole

The rapid extermination of sea cows was caused by thoughtless,
rapacious forms of exploitation which led to the unjustified annihilation
of a great number of animals. As seen from the descriptions of the
methods of hunting, success was only ensured in cases where many
people at once (not less than 20) were hunting one animal. Attempts to
get an animal by a small group of hunters were often useless. The
above-mentioned P. Yakovlev writes of this as follows: “. . . those
hunters live in yurts or winter houses in different places along the
northern shore of the Commander Islands in groups of 2 and 3 hunters
and they hunt for arctic foxes and other marine animals. They do not
have any other food except the meat of [sea] cows; the herds of cows
live in the sea near the shore and are frequently wounded by people,
but those wounded cows swim out to the sea and there become further
weakened by their wounds. They are later washed ashore after a long
period of time when their meat becomes bad and inedible.

By this sort of hunting sea cows are eradicated, and although many
are slaughtered (my emphasis, V.H.), not a single fresh sea cow comes
to their hand, and the hunters therefore suffered from starvation, and
the sea cows were exterminated.” Yakovlev foresaw the quick exter-
mination of the sea cows on Bering Island and tried to take measures
for the protection and wise use of the stocks of this most important
animal at that time. “For this purpose, on his (Yakovlev’s) return from
one sea voyage (i.e. sailing to the Commander Islands, V.H.) it was
reported on the 27th of November, 1765* to the Nizhne-Kamchatski1
Ostrog [fort] that hunting of the sea cows should be prohibited by order
(Dokladnaya Zapiska, V.H.) of the Bolsheretsk office in order to save
one Commander Island [Bering] from devastation because Mednyi
Island had already been devastated . . .” (Pekarskii, 1867). However,
as is known, nothing came of Yakoylev’s attempt. In this way, the sea

expedition consisted of 33 persons, could not spend the winter on this island in 1754,**
1.е., 13 years after the discovery of sea cows on Bering Island, because sea cows were
absent... . “Mednyi Island was found empty—he wrote—and ... for the expeditionary
force to spend the winter was impossible—due to the absence of sea cows the inhab-
itants of this island suffered from hunger.” Further, he wrote that “before the sea cows
were found in large herds but now these cows were destroyed, so none of them could
be seen, and now in the Mednyi Island there is not a single sea cow ...” (Pekarskii,
1867). Yakovlev and his “command” spent the winter on Bering (“Commander”) Island
(see below).

*This date should be 1755, since 13 years later, in 1768, the last one was killed
on Bering Island (see previous page)—D.D.

**Misprinted 1745 in Russian original—D.D.

cow at Mednyi Island was destroyed ten or twelve years after
its discovery. After approximately the same period, they stood under
threat of extermination at Bering Island, and 13 years following
Yakovlev’s attempt to protect this animal, was also completely
destroyed there.*

All information about the existence of the sea cow, which occa-
sionally appeared in the literature at the end of the eighteenth and the
beginning of the nineteenth century, as was shown by K. [von] Baer
(1838) and J.F. Brandt (1846), cannot be credited. Separate informa-
tion appeared later concerning some of these animals (for some, see
Grekov, 1958). The most famous of such reports, which seems most
plausible, is that of Nordenskidld (1882), in which it was mentioned
that a sea cow was observed on Bering Island in 1854. As was ex-
plained by Stejneger (1883, 1885), who interviewed the same people
to whom Nordenskidld referred, this information was found to be false.
Stejneger (1883) suggested that these animals were whales of smaller
size or, probably, female narwhals, which very rarely passed so far to
the south and therefore were considered unusual for the Aleutians.
Some other unreliable and unconfirmed information is not likely at all
and is not worth discussing. Some of it is simply fantastic (observations
in. . . California!).

Even the slightest probability is lacking in the latest information
given by Berzin, Tikhomirov and Troinin (1963), on the observation of
a sea cow in July 1962 at Cape Navarin (!), i.e. somewhat north of 62°
N. Lat. This information contradicts all the known facts about this
species, which are partially given above. It was based on complete
ignorance of the question and its literature (even the new literature), on
the biology of the animal, characteristics of its habitat, and its behavior
as well as the history of discovery and exploitation of northeastern
Siberia and Russian America in the eighteenth and nineteenth centuries.
This was actually the result of naive pursuit of sensation.

Still, in our time, remains of sea cows are sometimes found. These
remains have become rare, although a skull was found in the decade of
the 30’s and two lower jaws in a well-preserved condition were col-
lected in 1960 (N.N. Kartashev, Zoological Museum of Moscow
University).

There are, in general, numerous bones and skeletons of sea cows
preserved in museums of our country and in foreign museums which

*See foot note on previous page—Sci. Ed.

were collected from the Commander Islands. It is interesting that at the
beginning of the last century, a masticating plate was the only material
at the disposal of [von] Baer, and Brandt (1846) had only one imperfect
skull. At the beginning of the 80’s, Stejneger (1883) collected more
than 11 skulls and believed that more skulls could be collected. B.
Dybovskii brought some specimens from the Commanders in those
years. Now, such collections are impossible* (V.H.).

*Specimens have been collected as recently as 1992—D.D.

tes ELE

ORDER OF CARNIVORES
Ordo CARNIVORA Bowdich, 1821

Superorder of Carnivores
Superordo FERAE Linnaeus, 1758

Order of Carnivores
Ordo CARNIVORA Bowdich, 1821

49 The order includes species of relatively generalized type and clearly,
sometimes very sharply, specialized ones. A few are large, the majority
are of middle and small size, while some are of very small size. They
are typically terrestrial animals, in few cases, semiaquatic, and as an
exception aquatic animals of various construction and appearance, from
heavy and clumsy animals to light, slender and very mobile. The tail
is usually long (about half the body length or more), rarely short, of
approximately the length of the hind foot.

The extremities are plantigrade, semidigitigrade or digitigrade, digits
are usually 5, but in some one is absent. Digits are provided with
claws, in some species retractile; the terminal phalanges are not flat-
tened and not widened, but always laterally compressed. One digit is
never opposed to the others. In the wrist, the scaphoid (scaphoideum),
semilunar (lunatum) and central (centrale) bones are fused. The articu-
lating surface of the astragalus is pressed inward. The ulna and fibula
are normally developed and separate (not fused).* The foramen
entepicondyloideum on the humerus is either developed or absent; the
third trochanter of the femur is absent. The clavicle is either absent or
rudimentary.

There are no more than 22 lumbothoracic vertebrae, usually 13
thoracic, and 7 lumbar. The vertebral column is, in the majority of
cases, very flexible. The number of caudal vertebrae varies greatly.

The form of the skull is extremely varied—in some animals it is
elongated with highly developed facial part and a relatively short cra-
nial one, while in others the skull is very short, with rounded outline
and with relatively large and voluminous cranium. All the transitional
forms are also present. The hard palate is continuous, the eye socket is
usually open, and only exceptionally superficially separated, but in the

*Sic; refers to radius-ulna and tibia-fibula being distinct bones — Sci Ed.

majority, it communicates widely with the temporal fossa. The eye
sockets are directed forward and in several, strongly approximate
(binocular vision). The tympanic bulla (with one exception—WNandinia)
is ossified. The wing-shaped (ectopterygoid) fossa (f. pterygoidea, s.
ectopterygoidea) is not manifested in the majority of cases due to the
absence of the corresponding independent process of alisphenoid.

In the greater number of species, the crests and zygomatic arches
are strong and widely separated, a fact associated with the great devel-
opment of the jaw musculature. Ethmoturbinal (ecto- and endoturbinals)
olfactory folds are 5—7; the maxilloturbinal has double flexures or branch-
ing. The articular condyle of the lower jaw is semi-cylindrical and the
glenoid fossa on the skull is elongated, occupying a transverse position.
It is usually more or less deep, sometimes so much so that its anterior
and posterior edges tightly envelop the condyle of the lower jaw. The
angular process of the lower jaw is small and with no inward flexure.

The dental system is heterodont and diphyodont. The complete

33) dee An 4
formula is а. ит = 48, but usually the teeth are much fewer.

The reduction occurs at the expense of the cheek teeth and mainly at
the expense of the posterior molars and (to a lesser degree) of the
anterior premolars. The number of incisors is only rarely reduced to

В 2
я (Kamchatka “beaver” or sea otter,* Enhydra) or to 3 (sloth bear,

Melursus). The minimum dental formula is ICP м. = 28 (manul,

Felis manul) and even СРМ. = 26 (fossil sabre-tooth tiger,

Smilodon, of Machairodontinae).
In one case (bat-eared fox,** Otocyon) the dental formula may be

[2 GER Mo == 46" 48 500
та а 5

*In Russian, Kalan—Sci. Ed.

**In Russian, long-eared fox—Sci. Ed.

'The greatest number among recent placentals. Probably, this increase is a
secondary character and is connected with the transition to a considerable extent to
entomophagy (mainly grasshoppers).

The teeth always are rooted, brachyodont, cusped-cutting (tuberculo-
sectorial), quadri- or multituberculate with sharp, or rarely blunt, apices.
The teeth are strongly differentiated. Incisors are small and the canine
is usually strongly developed. The fourth upper premolar and the first
lower molar are usually large, with sharp cutting apices (“carnassial
teeth”), rarely with a more or less flat, bluntly tuberculated, mastica-
tory surface. A true diastema is absent.

The stomach is simple; the caecum is normally or weakly devel-
oped or absent. The brain is macrosmotic, the hemispheres are large
with three suprasylvan fissures. The pelage is always well developed,
composed of different hair categories (underfur, guard hair, directional
hairs), sometimes very dense, relatively long and fluffy, and sometimes
sparse and rough. The tail is usually fluffy. Vibrissae are well devel-
oped. The color varies greatly, being more or less monotone, or light
with bands, spots, shabrack*, etc.; some species are polymorphic. The
skin glands, with some exceptions, are normally developed; the major- .
ity of glandular complexes and large, complicated, often odiferous glands
(musky stink glands of skunks, etc.) develop in the anal region in
connection with the male genitalia. The milk glands—inguinal and ab-
dominal—range from 6—7 to 2 and one pairs. The scrotum is well
developed or else the testes are located subintegumentarily, but always
outside the abdominal cavity (“extra-abdominal”), and behind the copu-
latory organ. In the latter (except in hyena) a bone (0$ penis, baculum**)
is present. The uterus is doubled or with 2 horns. The placenta is
deciduous and zonal’.

Sexual dimorphism is absent or weakly manifested except in the
general size of the species with rare exceptions (lion). Age dimorphism
in some species is quite significant (wolf, lion and others), while in the
rest it is not manifested. Seasonal dimorphism, mainly in the density
and length of fur in the forms of temperate and northern latitudes, is
quite great and sometimes connected with changes in color, in which
case it is sharp (arctic fox, ermine), while in the others it is weak or
absent. In the forms of temperate and cold latitudes that are active in
winter, there are two molts which usually occur rapidly. In species of
those districts which spend the winter sleeping, the single molt extends

*Lit., goatskin; refers to a goatskin saddlecloth, hence by extension, to a “saddle”
pattern on the back—Sci. Ed.

**Spelled bacculum in Russian original—Sci. Ed.

Occasional reference to the placenta as of the discoid type (for example, Novikov,
1956) is a mistake. This was, apparently, based on a slip of pen in the diagnosis of the
order by Weber (1928).

over almost the whole summer (the summer fur is considered a tran-
sient coat). In warm countries, there is usually one molt.

Carnivores differ greatly in size—the largest has a head-body length
(excluding tail) to 300 cm and a weight to 725 kg and even | metric
ton (polar bear). The smallest form—the least weasel (Mustela nivalis
pygmaea) has a body length of 115—140 mm and a weight of about
100 g; i.e. the smallest weight is less than the greatest one by about
10,000 times (in case of artiodactyls this proportion is about 1 : 1,150).

In general constitution, proportions and external appearance, bio-
logical types, adaptation forms and other characteristics, the carnivores
are one of the most variable if not the most variable, of the orders of
mammals, including forms that are comparatively generalized types
(some bears) as well as extremely specialized ones (Kamchatka bea-
ver, or sea otter).

Among the carnivores there are heavy animals of low mobility and
slow movement (bears) as well as light and slender, rapid and swift
runners (wolf, cheetah), excellent swimming forms closely connected
with water (otters) and excellent climbers, and even partially special-
ized arboreal animals (some martens, viverrids and cats); some are
excellent diggers (badger), etc. Carnivores inhabit all latitudes, land-
scapes and the vertical montane belts. They live individually or in
pairs, and some are found in small packs (wolves, hunting dogs) at least
in part of the year. The majority are sedentary and are fairly strictly
restricted to their hunting areas; others wander widely, while several
accomplish regular seasonal migrations.

Some species are associated the entire year with holes (badger);
others, only in some parts of the year (time of reproduction) and a third
group has only open lairs even in the lactation period. Some are asso-
ciated continuously, or only at the time of reproduction, with tree
hollows, rock clefts, caverns and other similar shelters. Several species
of cold and temperate latitudes hibernate in winter. A series of species
are monogamous and live in pairs for more than one reproductive
season, while others form pairs only for one cycle and in others, the
male does not take any part in provisioning the young at all. True
polygamy (in the form of a “harem’’) does not occur.

The young range from 1—2 to 13 and even 20-22.’ However, the
number of young in the litter varies strongly within one species. The

3Maximal known number is in the wolf (13) and in arctic fox (22). Normally, these
numbers are much less.

offspring are born helpless, blind with closed ear openings and undi-
vided fingers, etc. and they develop and grow comparatively slowly.

Carnivores are extremely variable in their food habits. The major-
ity are sarcophagous, at times very narrowly specialized, feeding on
the meat of mammals and birds, and rarely ichthyophagous and polypha-
gous, in the diets of which a significant role is played by lower
vertebrates, terrestrial and aquatic (mainly fresh water) invertebrates
and plants. The number of species that are chiefly plant-eaters
(phytophages) is few; specialized entomophages comprise exceptions,
(aardwolf*—Proteles).

Among the sense organs, smell and hearing are well developed,
sight is usually weak, and not color. The external ear pinnae are very
different in size, from equal to head length (fennec) to almost undevel-
oped (otter).

Carnivores are very widely distributed, occupying nearly the whole
world. Aboriginal species are absent only in Australia and in some
islands and island groups. In the north, they are met with in the north-
ern most islands (New Siberian, Severnaya Zemlya, Franz Joseph Land,
Arctic archipelago of [North] America, Greenland) and even in the ice
of the Arctic Ocean, considerably to the north of these islands, all the
way to the pole (polar bear, arctic fox). The species of other orders,
even the pinnipeds, do not penetrate so far north. Carnivores are absent
on the island of the West Indies and the other islands of the Atlantic,
excépt for Newfoundland, Iceland and Jan Mayen. They are also ab-
sent, with the exception of the Falklands, on the islands of the southern
oceans, small islands of the Indian Ocean, in Tasmania, New Zealand
and New Guinea, Vetar [Wetar] and Ombai, in the Lesser Sundas (on
others, including Timor in the east, they are present), Tenimber
[Tanimbar], Kei and the majority of the Moluccas (except for
Halmahera, Buru and Ceram), in the Talaut [Talaud] archipelago and
on all the islands which lie to the east of New Guinea and on the
Hawaiian and Galapagos Islands. They are found in Ara, Sulawesi
(Celebes), Philippines, Ryukyu, Kuril, Sakhalin, Commander, Aleutian
and Pribilof islands.

The geographic range of the order has not essentially been changed
since historical time and it was not reduced—with some exceptions
[extirpation of the Falkland “wolf”, Canis (Dusicyon) australis
= antarcticus]. The geographic range was artificially widened for some

*In Russian original, maned “wolf’—Sci. Ed.

‘rouday ‘N'A ‘$8рон smjwjaundosnv snojuvanf sajsadsap] ‘QS00SUOU рэхцеиоэв OY} зэл| эти SAMA JO зризт э4з UC ‘(..о8шр 51и0),,)
oSuIp oy} Jo э8иел oY} зэоиэр в|ецзпу Зшрипоми$ sul] oY L ‘(рэзоплззиооэт) BIOATUIeD ‘sIoyepoid |е35эшэ} oy} Jo э8иег o1ydeIoeH [| ЗЫ ZS

091
Е Е
о
«ль “о о
"Е Te . Wy
>. eer
% ee
[.]

species; the Indian mongoose was established in some islands of the
West Indies, Fiji islands and several others; the European red fox was
introduced into Australia, etc. The dingo (“Canis familiaris dingo’),
occupying the entire continent of Australia, is not an aboriginal species.
This is a feral dog, which was introduced into Australia a very long
time ago by the ancestors of the contemporary Australians as a domes-
tic animal.*

On account of the diversity of biological types and forms of spe-
cialization which allow carnivores to occupy almost all habitable media,
the breadth of their geographic distribution, the diversity and abun-
dance of species, the relatively high numbers of many of them, and
other characters, carnivores without doubt may be considered a flour-
ishing and quite recent group. Although anthropological factors have
negatively influenced a series of species, decreasing their geographic
ranges and numbers, this influence is, however, generally less in rela-
tion to this order than if compared to several others. The “resistance”
of carnivores to this very strong destructive factor is an indication of
their vitality.

Concerning systematic relationships, carnivores are closely related

-to pinnipeds (Pinnipedia). The latter are considered to be derived di-
rectly from carnivores comparatively recently (Oligocene). All their
characteristics have been formed as sharp and direct progressive adap-
tations to an aquatic (marine) way of life. Some more primitive recent
pinnipeds (Otariidae—eared seals) are, in their least specialized charac-
ters (skull), very similar to the generalized forms of Carnivora (Steller’s
sea lion,* Eumetopias). Many authors unite carnivores and pinnipeds
in a single order Carnivora, dividing both groups only into the rank of
suborders: Fissipedia and Pinnipedia. Such a view was mainly based on
paleontological data. The alignment of the recent groups (by the “hori-
zontal” system), however, is such that for the purpose of keeping with
the sequence in the mammalian system, it is more natural to consider
Carnivora and Pinnipedia as separate orders.°

“The origin of the dingo from the primitive dogs of the tropical zone (Shensi type
“Canis familiaris schensi’) is indicated by the fact that the species of the genus Canis
are absent in the Malayan archipelago, and they (wolves and jackals) begin to occur only
in Indo-China and India.

*In Russian, sivuch—Sci. Ed.

Accordingly, in this edition, the suborder Fissipedia is opposed, not to the subor-
der Pinnipedia, but to the suborder Creodonta. In Simpson’s system (1945), Creodonta,
Fissipedia and Pinnipedia are considered as suborders of the order Carnivora.

The features that approximate carnivores with insectivores are
relatively great, and those which link them with artiodactyls and peris-
sodactyls are strongly divergent, and essentially in a purely historical
relationship (see vol. I). Their relationship with whales, which is fre-
quently emphasized and sometimes regarded as very close, is apparently
not more close than the relationship to ungulates.

The ancient and most primitive group of Carntvora—Creodonta—has
a great phylogenetic significance. They stand at the root of the exten-
sive cohort Ferungulata, comprising the orders of pinnipeds (Pinnipedia),
tubulidentates (Tubulidentata), probiscideans (Proboscidea), hyraxes
(Hyracoidea), sirenians (Sirenia), even- and odd-toed ungulates
(Artiodactyla, Perissodactyla) (see p. xx).

Carnivora is one of the most ancient orders of recent mammals,
stemming indirectly from the insectivores, the most primitive and an-
cient placentals. The primitive carnivores, the lower Creodonta
(Arctocyonidae), appear in lower Paleocene (North America) and are
represented by forms which are intimately connected—in certain re-
spects “transitional”’—with primitive Cretaceous insectivores
(Insectivora). Creodonts lived through their flowering in the Eocene
and were still relatively well, but notably less richly developed, in the
Oligocene. Their last representatives (Hyaenodontinae) survived only
to the very beginning of the Miocene. True Creodonta were represented
by several families (Arctocyonidae, Mesonychidae, Oxyaenidae,
Hyaenodontidae) which did not leave any trace. On the same level with
these families of the typical creodonts, the family Miacidae existed
from the middle Paleocene (became extinct in the Eocene), which by
general agreement, was the beginning of modern carnivores (Fissipedia).
The unique features of this family—which are, to a certain extent, of
“intermediate” character between Creodonta and Fissipedia—gave some
authors the basis for considering them among the Creodonta and gave
others the basis to relate them to the existing Carnivora and to ally
them to viverrids (Viverridae), or sometimes to cats (Felidae) or to
wolves (Canidae). If the Miacidae are considered among the existing
carnivores (Fissipedia), then the origin of this latter group (from the
primitive Creodonta) must be dated to the middle Paleocene. The re-
maining (recent) families of true carnivores (Fissipedia) appear not
earlier than the upper Eocene and the lower Oligocene.

Among the sum of characters, the contrast between the extinct and
recent groups of carnivores is relatively small, although in the extinct,
even a separate suborder is recognized (see below). It is less than, for

55)

example, in the “ungulate” group. The diversity of the group in the past
was similarly not greater than in the more recent (and present) time.
The same applies to the general number of species in different preced-
ing epochs. Therefore, in its historical aspects there is basis for
considering the order of carnivores as an existing flourishing group.

Classification of the order of carnivores, in spite of the abundance
and diversity of species, is, п: its main features, simple and does not
result in any serious disagreements. Two suborders are admitted—the
extinct Creodonta with four families (see above) and the recent
Fissipedia. The latter is divided into three superfamilies: the extinct
Miacoidae with one family, Miacidae; the recent Canoidea (Arctoidea
auct.) with four families—Canidae (wolves), Ursidae (bears),
Procyonidae (raccoons), Mustelidae (martens) and the recent Feloidea
(Aeluroidea, Herpestoidae auct.) with three families—Viverridae (viver-
rids), Hyaenidae (hyaenas), Felidae (cats).°

Canoidea are characterized by the following characters.
Tympanicum formed by the whole outer wall of auditory bulla. Parti-
tions of internal cavity of auditory bulla derived from cartilagenous
precursors, or precursors of the tympanicum. Canalis caroticus long.
Maxilloturbinalia large and branching, and partition off the nasoturbinalia
and ethmoturbinalia from the external nasal openings. Cowper’s glands
absent. Os penis large.

Feloidea are characterized by the following characters. Tympanicum
is, to some degree or other, semicircular and forms only the anterior
outer (tympanic) part of the auditory bulla, while the posterior inner
part is formed by the entotympanicum (os bullae). The tympanal-
entotympanal partition extends deep inside the bulla at the place of
divergence of both bones. Canalis caroticus short or absent.
Maxilloturbinalia small, doubly flexed, and do not partition off the
nasoturbinalia and ethmoturbinalia, which, in this way, approach the
external opening of the nasal passage. Cowper’s glands present. Os
penis small or absent.

Canoidea and Feloidea are sometimes considered to be suborders.

The systematics of carnivores is insufficiently worked out as re-
gards the genera and species. In particular, there is a strong tendency

°Attempts were often made to divide some families into several. Among Procyonidae
in particular, the [lesser] panda (Ai/urus—Ailuridae) and giant panda* (Ailuropoda—
Ailuropodidae) were divided into separate monotypic families. These, and also other
arrangements, cannot be considered well-founded and are not widely accepted.

*In Russian, bamboo bear—Sci. Ed.

here towards narrow interpretation of the genus (especially cats, bears

and some others) and species (particularly in America).’ According to

a more or less moderate opinion (Simpson, 1945), the order comprises
338 genera belonging to 12 families, of which 5 are extinct. Extinct

genera number 241; recent—97.®

Only a very approximate count can be made concerning the num-
ber of the recent species of carnivores at the present moment. Their
general number is approximately 240 (probably less), constituting only
8% of all existing mammals. On the average, the genus comprises 2.5
species. The greatest number of species is in the families Viverridae
(about 75) and Mustelidae (about 70); in Canidae there are 29, Felidae—
about 36, in Procyonidae—about 10, Ursidae—7, and Hyaenidae—4.
The greatest number of species live in Africa (about 100) and Eurasia
(about 90); in South America—about 50 species, in North America—
about 35.°

In practical relationships, carnivorous animals are one of the most
important groups of mammals. Among them, there are a great number
of fur-bearing species, some of the most valuable (sea otter, sable),
pests of livestock and hunting enterprises (wolf, raccoon dog, yellow-
throated marten), destroyers and regulators of a number of harmful
animals (mainly rodent—foxes, some martens, viverrids and others),
valuable sport-hunting animals (large species of cats, bears), vectors of
some domestic animal and human diseases (rabies), animals giving raw
material for perfumes (musk), etc.

Some species have been acclimatized in alien countries as fur-bear-
ing animals (raccoon dogs in European part of the USSR, American
raccoon, American mink in the USSR), as destroyers of harmful ani-
mals (snakes in the West Indies—mongooses). However, these measures
are not widely distributed in the sense of territory and number of species.
A series of species, as a result of the excessive utilization in connection
with the value of their fur or pursued for the sake of destruction for one
or another reason, sometimes very strongly, were reduced in number

"Lately, a wider approach to the species and genus concepts has been applied in
the systematics of carnivores, and it is well-known to be found in a revisionary stage
(see below in characters of families).

‘The relatively high “index of contemporaneity” of carnivores indicates that con-
temporary genera constitute a sufficiently large part of the total number of genera—more
than 28%.

*According to Anthony, 1928; Allen, 1939; Cabrera, 1957—1961; Chasen, 1940;
Carter, Hill and Tate, 1945; Anderson, 1946; Ellerman and Morrison-Scott, 1951;
Roberts, 1951; Haltenorth and Trense, 1956; Haltenorth, 1958 and other sources.

and distribution (sea otter, lion, tiger, bears, wolf, leopard, puma and
others). Some forms are exterminated completely or almost completely
(Falkland wolf, Cis-Caucasian leopard, Persian and Atlas lion and
others). A series of species find themselves under a complete or partial
[protective] ban, and some others are kept in preserves. The hunting of
the majority of fur-bearing and sport-hunting animals is regulated.
Conservation of carnivores usually gives good results.

Carnivores gave rise to several domestic animals. The most impor-
tant of these is the dog (“Canis familiaris”) which is the most ancient
of all the domestic animals, derived from the wolf (Canis lupus).'° The
domestic cat (“Felis catus”) is a minor, mainly decorative, domestic
animal, obtained as a result of the domestication of Felis sylvestris
libyca. The white polecat (Mustela putorius furo)}—the albinistic form
of the polecat—is only a semidomestic form from western Europe. In
the last decades, a series of species (red fox, Vulpes vulpes—imelanis-
tic form; Arctic fox, Alopex lagopus—normal and “blue” forms;
American mink, Mustela vison; raccoon, Procyon lotor; sable, Mustela
zibellina, and several others) became the objects of different forms of
fur farming, mainly hutch farming, and many different breeds of foxes
or minks have been derived.

In the fauna of USSR there are 41 autochthonous and 2 exotic
acclimatized species, i.e. about 17% of the species of the order, and
about 14% of the species of the USSR fauna. Both superfamilies, 5 out
of the 7 families (71%) and if the acclimatized American raccoon is
considered, 6 families (85%), and 22 genera out of the 97 existing ones
(22.5%) are present in the USSR."

The geographic range of carnivores in the USSR covers the whole
territory of the country and the Soviet sector of Arctic Ocean (they are
now exterminated in the southwestern part).

In the USSR, representatives of the order are found in all natural
conditions, including extremes of high mountains, deserts and the sea-
shore.

In the Soviet Union, carnivores constitute a very important group
practically. They include the most valuable fur-bearing animals, some
of them represent objects of hunting sports, while certain species play
a more or less considerable role as pests of livestock and hunting

‘The presumption of the participation of jackal, i.e. the diphyletic origin of the dog,
is to the highest degree improbable and is now entirely rejected.

''The number of genera, also mentioned above, is that given by Simpson (1945).
Later in the text, several genera will be discussed in a broader manner.

enterprises, carriers of some diseases, and destroyers and regulators of
the numbers of harmful rodents. The hunting of almost all species is
regulated (definite times of hunting, licensed trade); some are found
under complete or partial protection, while certain species are com-
pletely exterminated from the country or from some parts of it (V.H.).

Key for Identification of Families of the Order of
Carnivores

(Identification by External Features)

). Five digits on hind foot.
). Tail short (shorter than the hind foot) and hidden in body fur
or hardly protrudes from it. Size very large, body length of
adult 1.4—2 т ог more..... Bear family, Ursidae (р. 586).
3 ( 2). Tail long (longer than hind foot) and always clearly protrud-
ing from the fur. Body length less than 1.2 m.

4 ( 5). Tail distinctly bicolored—6—7 dark transverse rings distinct
against ‘lighter backeround.!2% jo. 44) hee ee Es
EUAN WUE LOTS LY Raccoon family, Procyonidae.*

5 ( 4). Tail one color, or terminal part darker than base, but trans-

verse, darkinings abSemts yin, (lity ahi 0h Veen abal core eee

6 ( 1). Four digits on hind foot.

7 ( 8). Four digits on fore foot. Along neck and back, high, protrud-
ing, shaggy crest of long elastic hairs. Color gray (without
reddish brown tones), with black stripes across the trunk... .
PARC Eh, AT я Hyaena family, Hyaenidae’’.

8 ( 7). Five digits on fore foot (first located somewhat higher than
remainder). High crest along neck and back absent. Color
different; if black transverse stripes on trunk, reddish brown
tones well-developed.

9 (10). Head elongated, with long dog- or fox-like muzzle. Tail
always reaches tarsal joint; fluffy, but hairs short at base,
thus tail is thinner at base than in middle. Claws not retrac-
tile, their tips directed forward and downward, slightly
compressed laterally and relatively blunt. Color without dis-
tinct spots and transverse stripes..................
о а ORES GENE lhe ai Wolf family, Canidae (p. 66).

*Described in Part 16.
"Described in the following volume.

10 ( 9).

D3).

NAN

Lh WwW
— —
<i

5 ( 4).

6 (1).

(8):

8 ( 9).
9 (10).

58 10 ( 9).

Head round with short, cat-like muzzle. Tail reaches the
tarsal joint, or is shorter, covered with hairs of equal length;
at base, not thinner than in middle. Claws retractile, their tips
directed forward and upward, strongly compressed laterally,
and very sharp. If claws not retractile, then of large size
(large dogs) and the color is light with small distinct, irregu-
lar black spots. Color with spots or transverse stripes, or
UMICOlOr hy. aes Е. ee ee Cat family, Felidae.”

Identification by Skull

Palatine bones protrude backward behind transverse line be-
tween posterior points of last teeth, and less than half [palatine]
width lies between these teeth’ (Fig. 12).

Four teeth in lower jaw on each side behind canine, five in
upper: totaltnumberzof teeth*s4 "7 What G2 WEL eel
Sid DG OLED GEE OP hele clea Hyaena family, Hyaenidae.”
Dental formula different.

Three teeth on each side in lower jaw behind canine; 4 or 3
in upper; total number of teeth 28 ог 30.............
Е: Cat family, Felidae."
Seven or 6 teeth behind canine in lower jaw, 6 in upper; total
number of teeth 42 or 40..... Wolf family, Canidae (p. 66).
Palatine bones protrude behind transverse line between pos-
terior points of last teeth; more than half their width lies
between these teeth'* (Fig. 12).

Size of skull large. Maximum length more than 200 mm....
ВА Mora te Samia cag Re Ui. Bear family, Ursidae (p. 586).
Size of skull smaller. Maximum length less than 200 mm.

Six teeth behind canine in upper jaw and 6 in lower; total
number of teeth 40..... Raccoon family, Procyonidae.*
Less than 6 teeth in upper jaw behind canine; 4, 5 or 6 in
lower; total number of teeth 38 or less..............
Я rece ae Marten family, Mustelidae.* (V.H.).

'3Described in the following volume.
'4Measurement taken along suture between palatine bones.
*Described in Part 1b.

Fig. 12. Skull of fox, Vulpes vulpes L., and wolverine Gulo gulo L., on right
(schematic). Characteristics of palatal construction and relation of the posterior
part of tooth row and palate. Sketch by N.N. Kondakov.

Superfamily of Wolves
Superfamilia CANOIDEA Simpson, 1931

Family of Wolves
Familia CANIDAE Gray, 1821

Carnivores of a more or less generalized type, of medium size, with
unusually long legs, light, well built and highly mobile.

Digitigrade; tarsal and carpal regions never touch the ground dur-
ing walking. On the anterior extremities there are five (in one case 4;
Lycaon) digits, of which I is always rudimentary and located consid-
erably higher than the others; on the posterior extremities, four digits.
The two middle digits are considerably longer than the rest. The claws
are not retractile, relatively blunt and not sharply compressed laterally.
The last phalanges are not capable of flexing far back and upwards and
have normal articulating surface.

The skull with long facial portion; small only in exceptional spe-
cies; swollen auditory bulla, with a more or less horizontal internal
partition, and the bone of the auditory meatus is of moderate length.
The lateral occipital (paroccipital) process is small, somewhat laterally

compressed, and adheres to the auditory bulla. The mastoid process is
small. The alisphenoid canal is well developed.
The dental system is relatively complete; the usual dental formula

is: Nas ye a), In one species it is [6 B= Me 38
ie mma Ио 2
44
it is: 1-С-Р-М-——=46-—48—50
(Speothos') and in another, it is: aa ees
(Otocyon).

The carnassial teeth are well developed, whereas the molars are
short and quite flat with low blunt cusps; in the upper ones, their width
exceeds their length. They have a grinding character, more or less
“omnivorous”. The canines are highly developed and are sometimes
very long and thin.

The caecum is fairly large ‘and spirally coiled or very weakly
developed and hardly visible. The glans penis is long and smooth
(without spines). An os penis is present, straight and bears a groove on
its upper surface.

The head is usually elongated with a long tapering muzzle. The
ears usually have a sharp apex and are located fairly high, of moderate
length, sometimes very long (longer than the head) and, as an excep-
tion, they may be short with rounded apex. The tail is of moderate
length (to tarsal joint) or longer, rarely short, and is covered all along
its length with long hairs (the hairs are shorter at the base). Cutaneous
glands are weakly developed, the anal glands are few, sometimes ab-
sent, and there are no genital glands; in some species, there is a scent
gland оп the upper side of the tail (“violaceous” gland). A scrotum is
present. There are 3-4-5 pairs of teats.

The pelage of northern forms, in winter fur, is long, dense, fluffy
and frequently silky, whereas in the south it is usually more coarse and
sparse. The general color is usually more or less monotone, gray,
yellow and red tones, sometimes with a dark dorsal saddle; and as an
exception, almost black or of large and bright irregular black, white
and red spots. In some species there are several color phases.

There is no sexual dimorphism; age dimorphism in some species is
rare in some or is quite sharply distinct. Seasonal dimorphism in forms
of temperate and northern latitudes is considerable, manifested in the

‘According to some data, this species has 36-38—40 teeth (Haltenorth, 1958).

density and length of fur, and sometimes, in color (arctic fox). There
are 1—2 molts per year.

In general constitution, species of the family are homogeneous,
representing a variation of one type as a whole—the wolf type. The
change in appearance is mainly determined by the proportion of the
trunk length and that of legs, and in part, tail length.

The form with the most extremely long legs, resembling the borzoi
dog, with a proportion of trunk length to shoulder height of about | : 1,
is represented by the South American maned wolf (Chrysocyon
brachyurus). The hunting dog of Africa (Lycaon pictus) is very close
to it. The wolf is of the same type, but with shorter legs. The subse-
quent type is represented by our fox and related species, which are of
still shorter legs and elongated trunk, with proportion equal to | : 0.5;
they have a long tail. The last group is represented by species which
are similar to the raccoon dog (Nyctereutes procyonoides), with the
extreme in this form represented by the South American bush dog
(Speothos venaticus). It has a somewhat “dachshund-like” appearance,
with proportions of 1 : 0.3 or slightly more, and a short tail.

The species differ considerably in size. The largest species—wolf
(Canis lupus)—has, in its large forms, body length to 160 cm, tail to
50 cm, height at the shoulders to 85 (100) cm and weight to 50-60 kg
or sometimes, exceptionally, to 80 kg. The smallest form—fennec
(Fennecus zerda)—has a body length of about 40 cm, tail of about 20
cm, height at the shoulders is about 20 cm and weight of about 2 kg.
The proportions of extreme forms by weight is, in this way, about
1 : 25-30.

The wolf family in its full breadth of specialization, both as a
whole and also in particular species, represents one of the relatively
“generalized” groups of the order.

The majority of the species of the family are excellent runners,
usually catching their live prey in open pursuit; some of them are very
quick and have extremely great endurance in running. They are not
adapted to climbing trees, and only the gray fox of America (Urocyon)
can climb somewhat. Some dig quite well. They live in all landscapes,
up to extreme high montane, but the majority, in all continents, are
connected with the more or less open areas (forest steppe, steppe,
desert). They frequently live individually, some live in pairs, and some
species apart from the breeding season live and hunt in family groups
or packs which are sometimes of great significance (hunting dog, Lycaon
pictus; red wolf [dhole], Cuon alpinus).

The majority are sedentary, but some species accomplish consid-
erable seasonal or irregular migrations or lead a nomadic life. They are
monogamous animals, some species live in pairs for many years. The
number of offspring [per litter] is up to 13—18 (according to some data
up to 22-23), but is usually 4~7 and not more than 10. Males of the
majority of species take part in the rearing of their offspring. This
family is psychologically one of the most highly developed groups of
carnivores, with definitely amicable internal relationships in packs and
families, hierarchies, etc. This is, to a considerable degree, associated
with the life in packs of many of the species.

They are principally nocturnal, frequently crepuscular and rarely
diurnal. Vision is relatively weak, hearing very good, and the sense of
smell is sharp to the highest degree. The fresh meat of small and
middle-sized mammals constitutes their main food, but several feed on
large ungulates, carrion and rarely insects and other invertebrates. They
utilize plant food, mainly fruits and berries, as supplementary food.

The geographic distribution of the family is exceptionally vast—
practically almost cosmopolitan. It corresponds to the geographic range
of the order; however, the species of this family are absent from
Madagascar, the entire Malayian archipelago with the exception of
Sumatra and Java, the Philippines, Hainan, Taiwan and the Ryukyu
islands. Occurrence in Australia (dingo) is secondary as a result of
domestic dogs becoming feral (see characteristics of the order). In the
north, the representatives of the family inhabit all lands and are found
on oceanic ice nearly to the pole (arctic fox). On the Falkland Islands,
they are extinct; several species (wolf) had their ranges reduced; how-
ever, the geographic area of the family as a whole did not essentially
change since historical time.

The family represents an ecological quite plastic and quickly adapt-
ing flourishing group of carnivores. The decline in numbers as well as
range reduction of some species occurs only as a result of excessive
hunting and destruction. Besides, the wolf, for example, persists even
in places with special hunts for its total destruction.

According to its systematic characteristics, the family occupies a
quite distinct position and is sharply distinguished from other families.
It is considered to be closest to the bear family, Ursidae. However,
judging by some characteristics of the genus Nyctereutes (raccoon
dog), the relation of this family to that of raccoons (Procyonidae) may
be closer (Frechkop, 1959). The resemblance of some forms to the

hyaena, Hyaenidae, is purely external, and is, to a certain extent, con-
vergent.

Concerning their origin, Canidae is one of the most ancient branches
of carnivores, already identifiable in the upper Eocene. The earliest
forms (Procynodictis, Cynodictis) are very primitive and possess fea-
tures indicating their relation to forms which lie in the root of all
Carnivora = Fissipedia.

Concerning the internal systematic structure of the family, it. is
quite unified. Attempts were made to divide it into some families by
separating such forms as the red wolf (Cuon) and hunting dog (Lycaon),
bush dog (Speothos) and, mainly, the bat-eared fox (Otocyon); how-
ever, these attempts were poorly based and did not meet with acceptance.

The present division into subfamilies is fairly simple and 1$ actu-
ally generally applicable. There are usually six subfamilies: the recent
Caninae (wolves, foxes, raccoon dogs), Simocyoninae (Cuoninae,
Lycaoninae—red wolves and others), Otocyoninae (bat-eared foxes)
and three extinct ones: Amphicyoninae, Amphicyonodontinae and
Borophaginae (Simpson, 1945). Sometimes, the genus Speothos (South
American bush dog) which, in the above-mentioned system is related
to Simocyoninae, is considered as a separate family—Speothoninae.
Among the existing forms, this species is so peculiar that this point of
view, supported most recently (Haltenorth, 1958), is represented as
most justified.

However, this system—at least that concerning the recent forms—
is not irreproachable. If the justified separation of the genus Otocyon
with its extremely peculiar dental formula in a separate group is ap-
proved (see above), this cannot be applied in relation to Simocyoninae.
The genus Lycaon (hunting dogs) of this group has the same dental
formula as all other Caninae (42) but it has only four digits on the fore-
limbs (the metacarpal bone of the first digit is even reduced); genus
Cuon (red wolf) has the same number of digits on the forelimbs as in
Caninae, but the dental formula is somewhat reduced (40, see below).
The differences in the skull structure of the members of these genera
from that of all Caninae in general, are not more than the differences
between the various forms of Caninae. The differences in the structure
of MI (see below) are also not very considerable and have no principal
significance. According to what has been previously mentioned, there
is no sharp limitation between Simocyoninae and Caninae in the char-
acteristics mentioned.

‘JoujdoH “D'A
‘(08uIp s1uDD,,) OBUIp oy} Jo эвиел oY} $э}оиэр епецзпу Зшриполи$ our] ‘эершео ‘soajom Jo Апиле} oy} Jo oduel 94413098 pajonysuosey “¢] “SI 19

4
Ц
\

‘2

is
`

The genus Speothos is better differentiated, first of all, by its
dental formula (38, see above), which is still more reduced than that
of Cuon, and by its “dachshund-like” appearance, blunt head, rounded
ears, short tail, etc. This form is sharply different, not only from Caninae
and Simocyoninae (Cuon, Lycaon), but also from the genera with which
it is frequently included in one group. Combining the characteristics in
the family allows us to think that it is more natural to divide this family
into two subfamilies—Otocyoninae (Огосуоп) and Caninae (all other
Canidae)—or to three, but in a combination of form different from the
adopted one: with the retention of Cuon and Lycaon in the Caninae, but
with the separation of Speothos in a special subfamily—Speothoninae.’

The number of genera in the family estimated by various authors
is different. Lately, 68 genera are accepted, of which 56 are extinct and
12 existing. Of the latter, eight genera belong to Caninae (Nyctereutes,
Canis, Alopex, Vulpes, Fennecus, Urocyon, Dusicyon, Chrysocyon),
three to Simocyoninae (Cuon, Lycaon, Speothos) and one to Otocyoninae
(Otocyon) (Simpson, 1945). Some, mainly on account of the American
forms, employ a greater number of genera. The actual number 1$, ap-
parently, 7-8: Caninae—WNyctereutes, Canis, Chrysocyan;
Simocyoninae—Speothos, Lycaon, Cuon; Otocyoninae—Otocyon. The
other genera usually employed are only subgenera, mainly of the genus
Canis (see below, Haltenorth, 1958).* The total number of species in
the family which was considered not long ago to be fairly large (about
45), equals 29; namely, 23 species of Canis, Alopex and Vulpes—
without the dingo and, excepting further reallocations in generic
characteristics—simensis (subgenus Simenia), rueppelli, chama,
pallida, zerda (subgenus Fennecus), microtis, thous (subgenus
Lycalopex), cinereoargenteus, culpaeus, hagenbecki, australis
(Dusicyon), Chrysocyon—| (С. brachyurus—South American maned
wolf), Nyctereutes—l| (see below), Speothos—1 (5. venaticus—South
American bush dog), Lycaon—1 (L. pictus—African hunting dog),
Cuon—1 (red wolves; see below), Оюсуоп—1 (О. megalotis—Afri-
can bat-eared fox).

*Systematic relationships in the family require special analysis, and here the
traditional grouping of the species of our fauna into two subfamilies is conditionally
given.

>The uniting of the genera Cuon and Lycaon, even without the application of
subgeneric rank, suggested by the same author, is not accepted. It is possible that
their union was based on the fact that the author wrongly applies the same dental
formula for both genera (42), although in Cuon there are only 40 teeth.

Six species inhabit Asia (V. сапа, Г. ferrilata, V. bengalensis*, Г.
corsac, N. procyonoides, C. alpinus); in Africa—seven (C. simensis,
V.** chama, V.** pallidus, C. adustus, C. mesomelas, L. pictus, O.
megalotis); in South America—seven (D.***. microtis, D.*** thous, О.
culpaeus, D. hagenbecki, D. australis, C. brachyurus, S. venaticus);
in North America—two (V. velox, C. latrans). One species lives in
Eurasia, Africa and North America (V. vulpes), two species in Africa
and Asia (V.** rueppelli, F.** zerda), two in Eurasia and North America
(C. lupus, A. lagopus), one—in Africa and Eurasia (C. aureus) and
one—in North and South America (D. cinereoargenteus).

Among the species of the family there are exploited animals which
give fur, usually expensive, and—to a lesser degree—are objects for
sport hunting. Some species are important destroyers of rodents. A
considerable number of them have no noticeable utility and, in general,
any practical significance. Some species are dangerous pests for live-
stock breeding and the hunting economy, and some are reservoirs for
rabies virus, transmitting it to domestic animals and man.

One of the species (wolf) has its geographic range considerably
decreased in connection with its eradication in many places both in the
Old and New World, and another one, Falkland wolf, Canis (Dusicyon)
australis, is completely extinct. The geographic range of some species
has increased in connection with their acclimatization (red fox). One
species is present in a wild form as well as in a completely domestic
form (wolf-dog) and two species are present in a wild and semi-domes-
tic form as objects of fur-farming (red fox and polar fox).

In the USSR fauna, 2 of the 3 subfamilies are represented: Caninae
and Simocyoninae; 5 of the 12 existing genera (42%)*: Nyctereutes,
Canis, Alopex, Vulpes, Cuon, and 8 species of 29 (about 27% of the
species of the family); they constitute about 2.6% of the total number
of species of our fauna.

The geographic range of this family covers the whole territory of
the USSR and its species are found under all natural conditions, to
extreme high montane regions and the ice of the Arctic Ocean.

*Misspelled /engalensis in Russian original—Sci. Ed.

**Spelled F.*probably for subgenus Fennecus, in Russian original—Sci. Ed.
***Spelled L., probably for subgenus Lycalopex, in Russian original—Sci. Ed.
The treatment of genera and subgenera is inconsistent—Sci. Ed.

4Or 3 out of 7, 1.е., 43%

The species of the family have an important value in the USSR as
fur-bearing and sport hunting animals, destroyers of rodents, pests for
animal breeding and hunting and carriers of some diseases (V.H.).

£2):

в,

314).

4 (5).

5 ( 8).

6 ( 7).

Key for Species Identification of the Wolf Family

Six teeth (premolars and molars) behind canine on lower jaw;
total number of teeth 40. Condylobasal length of skull not
less than 170 mm. General dimensions slightly less than those
of wolf. Color bright reddish-yellow; dorsal side of ear of
same color. On fore limbs, pads of middle digits connected
posteriorly by an anastomosis (Fig. 17).............
мои red wolf [46 ое], Сиоп alpinus (р. 571).
Seven teeth behind canine in lower jaw; total number of teeth
42.° Color different; if red, dorsal side of ear black.
Posterior lower angle of lower jaw below angular process
with sharp, deep angular groove (Fig. 14). Muzzle with dark,
mask-like shading, also covering eye region. Size similar to
р raccoon dog, Nyctereutes procyonoides (р. 82).
No sharp angular groove on posterior lower angle of lower
jaw, or only a very gently sloping, not deep arciform groove.
Dark “mask” on muzzle covering eye region absent.
Frontal (interorbital) region of skull noticeably elevated rela-
tive to nasal region, and line of upper profile of skull forms
protrusion here. Postorbital processes convex above. Apex of
lower canine, when jaws are closed, does not reach margin
of alveolus of upper canine. Facial part of skull relatively
short and broad (distance from posterior margin of preorbital
foramen to posterior margin of canine alveolus less than width
of skull above canines).

Tail with hair equal to or shorter than half of body length.
On undersurface of hind foot, five large bare pads (callosi-
ties), four digital and one behind them. Pupil rounded.
Arcuate line formed by anterior margins of nasal bones lacks
sloping middle process (near suture of both bones). Protrud-
ing cingulum on outer side of base of first upper molar (second
tooth from rear) narrow and blunt with median constriction or

‘As an individual difference, the number of molars and premolars may be 6.
However, in this case, the longest skull is 125 mm.

Fig. 14. Lower jaw of fox, Vulpes vulpes L., and raccoon dog, Nyctereutes
procyonoides Gray, below (schematic). The deep notch on the lower posterior
angle of the jaw of raccoon dogs is shown. Sketch by N.N. Kondakov.

gap. On fore limbs, pads of two middle digits not united by
commissure in posterior part (Figs. 15, 16, 17).
Dimensions large, skull length more than 200 mm; total
body length of adults more than 90 cm. Tail length almost
half of body length......... wolf, Canis lupus (p. 164).
7 ( 6). Arcuate line formed by anterior margins of nasal bones has
in middle (at suture of bones) small blunt and anterior pro-
jection. Cingulum on base of first upper molar wide, sharply
defined and without gaps. On fore limbs, pads of two middle
digits are connected by commissure in posterior part (Figs.
15, 165417):
Dimensions smaller, skull length not more than 190 mm;
total body length of adults less than 90 ст.б Tail length is
about half that of body ... . jackal, Canis aureus (р. 129).
8 (5). Frontal (interorbital) region of skull not noticeably elevated
relative to nasal region, and line of upper profile of skull
relatively uniformly elevated towards rear. Postorbital pro-
cesses flat or concave above. Apex of lower canine, when

бп the species, аз a rare exception, body length may be somewhat larger. In such
case, identification is decided on skull size, structural craniological characters, and
structure of the pads of the fore limbs.

64 Fig. 15. Anterior margin of nasal bones of wolf (to the left) and jackal—dorsal
view. Sketch by N.N. Kondakov.

65 Fig. 16. Cingulum of anterior upper molar of wolf (above) and jackal. Sketch by
N.N. Kondakov.

65 Fig. 17. Sole and digital pads of fore limb of wolf, jackal and red wolf (left
to right). Sketch by N.N. Kondakov.

9 (10).

10 ( 9).

№ (2).

12 (1).

13 (14).

idl

jaws are closed, reaches margin of alveolus of upper canine
or extends beyond. Facial part of skull relatively narrow and
long (distance from posterior margin of preorbital foramen to
posterior margin of canine alveolus more than or equal to
width of skull above canines).

Tail with hair exceeds half of body length. Undersurface

of hind foot has only 4 circular, bare digital pads (callosi-
ties). Pupil oval, vertical.
Postorbital processes above flat or slightly concave. Apex of
lower canine, when jaws are closed, reaches only margin of -
alveolus of upper canine or extends a bit farther above it
(Fig. 18). Distance from posterior margin of infraorbital
foramen to posterior margin of canine alveolus equal or
almost equal to skull width above canines.

Ear with rounded tip, relatively short; when bent for-
ward, tip does not reach eye. Body color either completely
white or evenly brownish-gray (winter fur), or back, limbs
and tail are brownish while side and venter are dirty white
BA Meola ihe, 159 AES PURE IAT arctic fox, Alopex lagopus (p. 274).
Postorbital processes above strongly concave (at base, dorsal
groove is formed). Apex of lower canine, when jaws are
closed, extends considerably past margin of upper canine
alveolus. Distance from posterior margin of infraorbital fora-
men to posterior margin of canine alveolus greater than width
of skull above canines.

Ear with sharp tip, long; when bent forward, tip reaches
eye. Color different.

Posterior side of ear brownish-black or pure black and sharply
distinguished from red or sandy-reddish color of back. Black
markings on anterior side of fore limbs. Extreme tip of tail,
at least very small part, white. Size relatively large; body
length more than 65 cm, total length of skull more than 115
cm, width of zygomatic arches more than 64 mm.......
BPs cant ро ahead DOM eR red fox, Vulpes vulpes (p. 469).
Posterior side of ear not black or brownish-black and not
distinguished from color of back. No black marks on anterior
side of fore limbs. Tail tip not white. Size smaller.

Back gray mixed with black (in form of ripples). Chin and
lower lip dark brown or black. Canines very long; when jaws
are closed, ends of upper canines protrude beyond level of

5
1 3 : 6
Fig. 18. Canines and incisors from side* (Sketch by М.М. Kondakov). 1—wolf;

2—jackal; 3—arctic fox; 4—Middle Russian red fox (Vulpes vulpes vulpes L.);
5—Turkmenian red fox (V. v. flavescens Gray); 6—Corsac fox.

lower margin of lower jaw. Size very small; body length less
than 50 cm, skull length less than 95 mm, zygomatic width
less than50mm...... Afghan fox, Vulpes cana (p. 390).
14 (13). Color of back is rusty or rusty-yellowish, grizzled from the
white ends of the guard hair. No mixture of black color. Chin
and lower lip white. Canines relatively shorter. Ends of upper
canines, when jaws are closed, reach level of lower margin
of lower jaw. Size larger; body length more than 50 cm, skull
length more than 95 mm, zygomatic width more than 58 mm
ии corsac fox, Vulpes corsac (р. 396) (У.Н.).

Subfamily of Wolves
Subfamilia Caninae Gill, 1872

Genus of Raccoon Dogs, or Manguts
Genus Nyctereutes Temminck, 1839

1839. Nyctereutes Temmink. In: Van der Hoevens Tijdschr. Nat. Ges.
Phys., 5, p. 285. Nyctereutes viverrinus Temminck.

Forms of small size.
Skull relatively quite heavy and massive, moderately elongated,
with narrow zygomatic** arches, and not high (height in occipital

*In Russian original, “front”, an apparent lapsus—Sci. Ed.
**auditory” in Russian original, an obvious /apsus—Sci. Ed.

region little more than one third of condylobasal length). Protuberances
and crests of skull well developed, sagittal crest high in old animals and
occipital quite considerable.

Facial region of skull short and its length nearly equal to cranial—
distance between anterior end of premaxillae and line connecting ends
of supraorbital processes equal or nearly equal to that between this line
and upper margin of foramen magnum. Facial region of skull massive.
Distance between posterior wall of infraorbital foramen and posterior
margin of canine alveolus less than width of skull above canines or
equal to it.

Interorbital (frontal) region elevated, somewhat swollen in region
of frontal sinuses between supraorbital processes; processes themselves
not large, flat or slightly concave above, with tip sharply recurved
downwards. Upper profile of skull at first almost straight and then
becomes convex arcuate line; at anterior of nasal region already abruptly
elevated and no frontal shelves occur. Posterior edge of tooth row does
not reach level of anterior border of interpterygoid notch. Hard palate
extends backwards behind line uniting posterior points of last molars
to approximately 1/3 the distance between inner corners of last molars.

At lower edge of lower jaw, anterior to angular process, a large,
massive protuberance, separated from angular process by deep angular
notch. Such a mandibular structure also present only in American gray
fox, Dusicyon (Urocyon) cinereoargenteus* (Fig. 14). Angular pro-
cess large and massive, with broad base and without upwardly directed
denticle.

Dental formula: ГРМ. = 42. As a fairly rare individual

deviation, there may occur i = 44. Teeth relatively small and weak.
3

Upper canines strong, but short, their ends far short of upper border of
lower jaw, and slightly curved; their ends not directed straight down-
wards, but somewhat slanted to the side. Cutting edge on posterior side
of canine and on lateral surfaces, a blunt, smooth vertical edge. Upper
carnassial teeth relatively weak. Incisors have no additional blades;
external [third] incisor not adjacent to second, and separated from it by
noticeable gap.

*Mispelled cinereoargentatus in Russian original—Sci. Ed.

Trunk long, legs short, and tail hangs below tarsal joint, but when
the animal is standing, does not reach ground. Ears of moderate length.

Pelage (in northern forms) dense and fluffy in winter. Molt occurs
once a year. Five pairs of teats. Intestine relatively long, compared to
body length being 4.3 : 1 (Korneev, 1954).

Inhabitants of closed landscapes—forests, river valleys, preferably
moist. Of low mobility, and travel relatively slowly. They do not per-
form regular migrations. Monogamous animals, pairing for one breeding
season. 10—15 offspring, usually less. Young born in burrows, and they
are associated with the burrows throughout the entire year. Apart from
the period of reproduction, they live singly. They are predators, feeding
on small vertebrates, especially rodents, amphibia, eggs, insects and
other invertebrates, and also on berries and fruits. In the northern part
of their range, they hibernate (the only species in the family).

The natural range of this genus is not great and is limited to part
of eastern Asia. It occupies the southern parts of the Amur [and] Ussuri
territories, northeastern China (former Manchuria), the Korean penin-
sula, eastern China to the extreme south, and the Japanese islands.

In the last ten years, the range has significantly expanded due to
acclimatization in almost the whole European part of the [Soviet] Union
and in the Caucasus, in separate places in southern Siberia and the
eastern part of Middle Asia (see below; description of species).

Because of its systematic characteristics the genus Nyctereutes
represents a strongly deviant form of the subfamily and stands com-
pletely apart. In any case the genus is more isolated from other genera
of the subfamily than they are from each other. Even by broad inter-
pretation of the genus, when such groups as Vulpes, Alopex, Fennecus,
Simenia, Lycalopex and Dusicyon are included in the genus Canis (see
characteristics of the family), the generic independence of Nyctereutes
remains obvious and undoubted.

The genus may be considered specialized in some respects; how-
ever the more basic features, mainly in the structure of the facial part
of the skull, in dentition, and in some other characters, are primitive.
The genus ought to be included among the least specialized in the
subfamily (family ?) and at the base of the systematic series. Also
primitive are such characteristics as poorly developed predatoriness,
low mobility (the animal is essentially not a predator, but a “collec-
tor”), large number of young and hibernation. Apparently, it is
sufficiently basic to consider the genus as archaic and relict. This is
supported by its geographic range, which is associated with a region in
which Tertiary relict forms are preserved.

seek

Fig. 19. Geographic range of the mangut, or raccoon dog , Nyctereutes Temm.
In the east, the natural species distribution of the mangut; in the west, the region
inhabited by acclimatized animals in the European part of the Soviet Union and
in the Caucasus. Arrows point to the direction of spread of the raccoon dog from
the USSR to Finland, Poland, Romania and Central Europe. (V.G. Heptner).

Nyctereutes has some characters similar to those of the raccoon
family, Procyonidae. Thus, both in general outlines and some structural
details, the skull of Nyctereutes resembles that of Bassariscus
(cacomistles), and the dentition occupies, in a certain sense, an inter-
mediate position between that of the true Canidae (Vulpes) and
Procyonidae (Procyon). The biological characters noted, especially

hibernation, are also “raccoon” features (Frechkop, 1959). These char-
acters do not provide a basis for including Nyctereutes in the raccoon
family, but they well emphasize the individuality of the genus. More-
over, they show that the relationship between the two named families
is closer than what is usually believed.

Fossil representatives of the genus are not known. However, it can
be assumed that the time of their development is concerned with the
Pliocene, and perhaps even earlier.

The genus included one species: Nyctereutes procyonoides (Gray,
1834),* i.e. about 3.4% of the species of the family. This species
constitutes about 0.3% of the species of the USSR fauna.

The natural range of the genus in the USSR is restricted to a small
strip of forests of the Manchurian type in the Ussuri and Amur terri-
tories. This species has been acclimatized in other parts of the USSR
(see below).

It is one of the secondary fur-bearing animals; useful to a certain
degree in destroying rodents, but it is essentially harmful to the wild
game economy (V.H.).

USSURI RACCOON, RACCOON DOG, MANGUT’
Nyctereutes procyonoides Gray, 1843

1843. Canis procyonoides Gray. Illustr. Zool., 2, pl. 1. Vic. Canton.

1907. Nyctereutes ussuriensis. Matschie, Wiss. Ergebn. Filchners. Exp.
nach China, 10, p. 178, Near Ussuri mouth.

1907. Nyctereutes amurensis. Matschie. Ibid., р. 179. Amur.

1958. Nyctereutes procionoides kalininensis. Sorokin. Byull. Mosk.
obshch. 'ispyt. prirody. Kalinin Department. 1. Kalinin district.
(V.H.).

Diagnosis
Only species of the genus.

*In Russian original “Temmink, 1839’—Sci. Ed.

’First and second names are artificial and literal. “Raccoon 402” is simply a
translation from the Latin—Canis procyonoides. It is the most common name in our
literature. “Ussurian raccoon” is a furrier’s term. The Russian local (Amur Cossack)
name is “enot” [raccoon] (Cherkasov, 1867). Mangut is an Evenk name. It is better than
the others, and was used by Russians. Unfortunately, it is not used in literature at the
present time.

83
Description

General appearance of the Ussuri raccoon is entirely unique, not resem-
bling the appearance of other representatives of the family. The trunk
is long, legs short and they look particularly short in the animals with
winter fur, which on the trunk is very long. Accordingly, the body has
a massive appearance. Also, the legs are covered with short hairs and
they look very thin. This gives the impression of a stocky and clumsy
animal on short, thin legs.

The fur in winter is not proportional to the size of the animal. It
is long (length of guard hair to 120 mm), thick, with dense underfur and
coarse guard hair, shaggy and stiff. The tail is relatively short—not
more than 1/3 the body length, its end falls below the tarsal joint, but
it does not reach the ground when the animal is standing. The hairs of
the tail are very long and dense, not shorter at the base than on any
other part of its whole length and the tail fur is not clearly distinct from,
and is transitional to, the fur of the croup. This reinforces the impres-
sion of a general elongation of the animal’s body. The head is not large,
with short, but pointed muzzle covered with short hairs. Behind the
eyes the hair rapidly increases in length and quantity, and quite large
ears, blunted on the ends, protrude slightly from the fur. There are large

Fig. 20. Mangut, or raccoon dog, Nyctereutes pracyonoides Gray. Sketch by
A.N. Komarov.

Fig. 21. Male mangut, 10-11 months of age. “Кедгоуауа Pad’ ” preserve
(southern Primor’e). March 1964. Photograph by A.G. Pankrat’ev.

“side-whiskers” of elongated hairs on the cheeks. The pointed muzzle
looks as if it is emerging from the long fur of the head.

The general color tone of the winter fur is dirty, earth-brown or
brownish-gray with a more or less considerable overlay of black (color
of guard hairs). The tail is considerably darker than the trunk. Along
the back extends a darker stripe which broadens on the shoulders,
forming there an unclear cross-shaped figure. The abdominal surface is
yellowish-brown and the chest is dark brown or blackish. Ears are
black posteriorly. In the eye region, in front of them and on the cheeks
below and behind the eyes and crossing the “side-whiskers”’, is located
a dark (almost black) field. Together, they form on the muzzle of the
mangut a characteristic picture in the form of a mask which contrasts
sharply with the brighter color of the muzzle and remaining parts of the
head. This picture in particular, and the general color tone gives the
described species a certain similarity to the American raccoon.

In summer, during the transition from one winter coat to another,
the mangut has at first (first half of summer) only a pelage of dry
lifeless top hairs (underfur sheds first) and later, the fur is formed of
short, incompletely grown guard hairs, almost without underfur. Such
hairs are several times shorter than winter ones. The color during that

time resembles that in winter, but its general tone is brighter, reddish-
straw, but with a mixture of blackish-brown. The dark color forms a
stripe along the back and dorsal side of neck, and a cross-shaped figure
in the shoulder region. The chest 1s brownish-black while the belly and
groin are brownish-gray with yellowish. The legs are blackish-brown,
the “mask” is well defined; the chin, neck and chest are black. On the
sides of the neck the color is quite bright, pale-rusty. Claws are light-
horn.

For description of skull, see description of the genus. The diploid
number of chromosomes is 42.*

Relatively speaking, data on the dimensions and weight of manguts
are extremely scanty. Body length is about 65—80 cm; tail length, 15—
25 cm; weight in summer 4—6 kg and in winter, 6-10 kg (С. Naumov
and Lavrov, 1948).

Dimensions of the skull (in mm) of males from the Far East are as
follows (42 specimens; Sorokin, 1958):

Range Mean

Greatest length 114.3—131.8 123.4 = 0.6
Condylobasal length 103.2—125.8 119.2 + 0.6
Zygomatic width 61.0— 75.0 67.2 + 0.4
Width of muzzle between 20.3— 24.9 223 + 0.1

canines
Width between orbits 19.8— 25.2 22.6 + 01
Distance between supraorbital

processes 26.0— 36.0 31.5: 03
Greatest width of skull 40.4— 44.5 ADS tee 01
Length of nasal bones 35.4— 50.7 42.6 + 0.4
Height of skull in region of

auditory bullae 41.3— 48.3 44.6 + 0.2
Length of upper tooth row 42.7— 49.4 46.4 + 0.2

Dimensions of the skull of females from the Far East
(40 specimens):

Range Mean
Greatest length 112.8—131.7 120.3. +, 2056
Condylobasal length (36) 110.1—126.6 109:8 ==, 0.5

*Now known to be 38 + 1-4 small В chromosomes Юг Japanese animals. Мат-
land raccon dogs have a diploid number of 54 + 1-3B chromosomes—Sci. Ed.

=
= ых у -- 12 SS
oR = ee LE
4 =“. с А
плели ТИТ = ae tT” =

В АИ
Ne a

ae?

. а а
АА
t

ог raccoon dog, Nyctereutes procyonoides Gray.

’

Fig. 22. Skull of mangut

Range Mean

Zygomatic width 60.2— 71.3 66.2 + 0.4
Width of muzzle between canines 18.7— 23.8 21.5 + 0.1
Width between orbits 19.5— 25.5 22.3 № 0.2
Greatest width of skull 38.9— 45.3 42.1 + 0.2
Length of nasal bones (39) 37.2— 49.7 43.6 + 0.4
Height of skull in region of

auditory bullae (39) 41.0— 47.6 44.6 + 0.2
Length of upper tooth row 43.2— 51.7 45.8 + 0.2

Difference in the dimensions of males and females is so small that
sexual dimorphism is basically not evident (V.H.).

Systematic Position
See characteristics of the genus.

Geographic Distribution

The forest areas of eastern Asia from the Amur territory to northern

Indochina. This is a typical range for a species of the Manchurian-
Chinese fauna.

As a result of acclimatization, a new range was established in the
European part of the [Soviet] Union and in the Caucasus, and there are
separate sections of occurrence in Siberia and Middle Asia.

Geographic Range in the Soviet Union

The range in the USSR represents the extreme northern border of the
range of the species. It is not large, and occupies a very small part of
the country; namely the Ussuri territory and the southern part of the
Amur.

The outlines of the range within the territory of the [Soviet] Union
are fairly complicated. In the east it begins at the Pacific coast of the
Ussuri territory in the region of the mouth of the Tumnin river (about
51° М. Lat.) and extends as a narrow strip along the seacoast to the
south and southeast along the foothills and lower mountain belt. Pass-
ing around the higher parts of the southern extremity of Sikhote-Alin’
from the south and approximately in the latitude of the northern end of
Lake Khanka, the boundary traverses the western slope of the range.
Along the Ussuri valley and along the western slopes of Sikhote-Alin’,

the range strip passes north to the Amur and reaches it at the mouth of
the Ussuri. Along the valleys of the more significant right tributaries
of the Ussuri (Bikin, Khor, Iman), the range at places extends far to
the east into the mountains; however, the mangut is absent in the
extreme high-montane Sikhote-Alin’, and in the region of the extensive
coni-ferous forests.

Along the Amur valley and adjacent places, the range extends
northward (downstream) nearly to the Khungara. From this place the
northern border of the range continues westward through the Burein
mountains and the middle course of the Bureya and Zeya to the Amur
in the region of the junction of the Shilka with the Argun. Because of
scanty information, it is impossible to determine whether the species
described is distributed all along the Amur valley to the west from the
Zeya, or whether they penetrate from the south (from China) to the
mouths of the Shilka and Argun. They are not found in the extreme
northern part of the northern bend of Amur. According to some data,
the northern border of the normal occurrence extends in our country
from the east only to the mouth of the Khumaerkhe on the Amur. To
the south, the range extends everywhere up to the state boundary.

The mangut is absent in Trans-Baikal, although in remote times, it
apparently reached the upper Onan.*® Sakhalin is not included in the
natural range.

In the last decades (starting from 1934), the Ussuri raccoon in-
creasingly has been established, especially in the European part of the
[Soviet] Union, and also transplanted to separate places in the Asiatic
part. At the present time in the European part of the country, the
mangut formed a vast distribution area, which considerably exceeds its
natural range within the USSR.

In the north the new range occupies all of Karelia and, probably,
the entire southern part of the Kola peninsula, the whole of Arkhangel’sk
district including the forest-tundra; however, incursions by the mangut
onto the tundra occur. Apparently, the northern border passes some-
what more to the south along the Pechora. In the west, the range in the
entire European part of the Union reaches the western state boundary
and in some places beyond it. In the south, the range reaches the Black,
Azov and Caspian seas. The eastern border is less defined. On the

$014 data about the permanent habitation of the animal here were not confirmed.
The range is according to Cherkasov, 1867; Radde, 1862; Maak, 1859; Schrenk, 1858;
Ognev, 1931; C. Naumov and Lavrov, 1941; material of N.N. Rukovskii and other
sources.

northeast, the mangut is known (settled) at the river Ishma (left tribu-
tary of the Pechora). To the south, the border extends, apparently,
along the Urals west of the range—through Perm district and Bashkiriya
to Orenburg. From the Urals it extends to the Volga and follows it
down to the [Caspian] Sea. The line of the eastern border of the
European range is very undefined, not only due to insufficient informa-
tion, but also because it changes as a result of introductions of the
animal (easternmost transplantations: Syktyvkar, Udmurtiya, Ufa,
Orenburg).

Within the outlined borders, by 1960 mangut actually occurred
everywhere. If it was still absent in more or less considerable parts
(some parts of Karelia and Cis-Urals) or its distribution was sporadic
or it was very rare (Arkhangel’sk district), then in most of the area
(several districts) its number increased immensely and it became an
important enemy of the hunting economy.

In the Caucasus, the Ussuri raccoon has been transplanted
and established in a series of places. Its range extends into the Cis-
Caucasian steppes and foothill districts, and the eastern half
of Trans-Caucasus. It is, apparently, absent only in the western part
of Georgia. Its occurrence within these outlined boundaries is not
continuous.

Outside the above-described borders, Ussuri raccoons were estab-
lished in Middle Asia and Kazakhstan—to the east of Frunze, in
Issyk-Kul, in the Dzhelalabad region, at the mouths of the Ili and
Lepsa, to the south of Alma-Ata, in the Dzhungarsk Alatau, and in
Katon-Karaga region in the southwest Altai.

In Siberia, manguts have been established at the mouths of the
Biya and Katun, and in a series of places in the Altai, near Tom’sk at
the sources of the Tom’, near Novosibirsk, near Krasnoyarsk, in Tuva,
at Irkutsk, northeast of Ulan-Uda and along the Dzhida and even in
Yakutiya—at the mouth of the Vilyui and southeast of Suntar on the
Vilyui. They were also acclimatized in several other places, including
Sakhalin.’ Establishment in Siberia and Middle Asia did not lead to the
formation of significant patches of occurrence, to say nothing of a
continuous range. In the majority of places, apparently, the animals
died or are dying, and in some places, small foci were formed.

*Acclimatization of the mangut from data of Lavrov, 1946; Rukovskii, 1950;
Korneev, 1954; Vereshchagin, 1959; Berens and Yakushevich, 1959; Marvin,
1959; and a number of other works as well as the materials provided by N.N.
Rukovskii.

08
0

>
>
=>
>=
=>
—
=

91
Geographic Range outside the Soviet Union

The range outside the USSR includes China, from former Manchuria
southward to Fukien, Guandong and Yunnan, the Korean peninsula, the
northeastern part of the Indochina peninsula (Tonkin, now Bakbo prov-
ince in North Vietnam). Borders of the distribution to the west are
undefined; in northeast China the range does not go beyond Great
Khingan (absent in the Mongolian Republic); to the south, it is known
from Shaanxi and Sichuan; absent in Burma. It is found in the Japanese
islands (Honshu, Shikoku, Kyushu, but is absent on the Ryukyus). It is
not present on Taiwan and Hainan.

As a result of the spread of the animals acclimatized in the Euro-
pean part of our country, at the end of the 1950’s the Ussuri raccoon
penetrated to Poland, Romania, Finland, northeast Sweden and even in
the G[erman] D[emocratic] R[epublic], where they are found east of
Berlin (V.H.).

Geographic Variation

Geographic variation of the species is not well studied. Five subspecies
are usually accepted. Within the Soviet Union one subspecies is present.

Ussuri mangut, N. p. ussuriensis Matschie, 1907 (syn. amurensis,
kalininensis).

For description, see above.

Ussuri and Amur territories. Acclimatized in the European part of
the USSR and in the Caucasus.

Outside the USSR, northeast China and Korean peninsula; the
limits of distribution to the south are unclear.

This form is poorly described, but by comparison with the nominal
form from the extreme south of the range it apparently is distinguished
by the somewhat larger size and denser, longer hair. There is no com-
parative study of our form. The name amurensis is based on an
occurrence character.

Fig. 23. Range of the mangut, Nyctereutes procyonoides Gray in USSR.
Continuous line in the Far East is the northern border of the natural range of the
species. The more or less continuous distribution formed as a result of
acclimatization in the European part of the country and in the Caucasus is shown
by the closed line in the West. Individual dots are the places of establishment of
the animals beyond the limits of the western range. Data of the western part of
the distribution area are from the beginning of the 60’s. (V.G. Heptner).

Separation of animals from Kalinin district in the form of an inde-
pendent subspecies М. р. kalininensis (Sorokin, 1959) was unnecessary
and uncalled for. As is evident from the figures given by the author,
there is no difference in the dimensions of the skull. Differences in the
quality of fur (longer, fluffier, softer and denser) are very difficult to
perceive. If differences exist, they mainly reveal completely normal
phenotypic variability. The independence of this form has neither a
practical nor theoretical basis.

Outside the boundaries of the USSR, the following forms are usu-
ally accepted: (1) N. p. procyonoides Gray, 1834—China, except
Yunnan, Tonkin; (2) N. p. viverrinus Temm., 1844—Japanese islands;
(3) N. p. koreensis Mori, 1922—Korean peninsula; (4) N. p. orestes
Thos., 1923—Yunnan. The reality of the last two forms, especially the
Korean one, is highly doubtful. Most probably there are 3 forms—the
mainland southern (procyonoides), northern (ussuriensis) and island
(viverrinus). (V.H.).

Biology

Population. There are hardly any data about the number of raccoon
dogs within the limits of its natural range in the Far East. The densest
population is observed in the southwestern part of Primorsk territory,
south of Lake Khanka, in the lower and middle courses of the Iman,
Bikin, and Khor rivers and the middle course of the Amur River. In the
basins of the Maikhe and Suchan rivers and along the coast of the
Japanese Sea south of Nakhodka Bay (Ussuriterritory) in 1931, the
population density (calculated by the number of burrows found) con-
sisted in various habitats of from 0.9 to 8 animals, averaging 3.4 per
1000 hectares. On Askol’d Island, density averaged 3.8 animals over
the same area [i.e., 1000 В] (Bannikov, 1964).

In the majority of regions of acclimatization of this species in the
European part of the USSR, in the zone of mixed and deciduous forests,
the population density constitutes about 1—2 animals per 1000 hectares.
In the flooded deciduous forests of some regions, for example Novgorod
(Morozov, 1953), it reaches 13 individuals per 1000 hectares; in the
middle Volga region and Tatariya, 2—3 animals (V. Popov, 1953), in
Gor’kovsk district, in different years, from 15 to 48 (Kozlov, 1952);
in Voronezh district an average of 1-2 animals per 1000 hectares, but
in some regions, for example Bogucharsk, to 20 individuals in such an
area (Obtemperanskii, 1953; Ivanova, 1962). In the northeastern

regions of Lithuania, 5 to 10 animals were counted in 1000 hectares of
forest area; in the northern regions, 2—4 individuals; and in the southern
and western regions, about 0.5 per 1000 hectares (Prusaite, 1961).

In Rostovsk district and in the Ukraine, the population density of
raccoon dogs consists of 2—5 animals per 1000 hectares of land. How-
ever, on the forested islands and along the shores of water bodies, it
reaches 29-46 animals (Rall’ and Kritskaya, 1953; Korneev, 1954;
Sergeenko, 1956 and others). In the northern Caucasus, in Dagestan,
Georgia and Armenia, Azerbaijan and Moldavia, the number of rac-
coon dogs, as a rule, does not exceed 1—2 individuals per 1000 hectares,
and frequently is much less (Stepanov, 1939; Dal’, 1941; Meladze,
1947; Kostyuchenko, 1950; Lavrov, 1957; Trushchalova, 1959; Aliev,
1962 and others). In Kazakhstan and Kirghizia, the density is even
lower but in small sections, for example, along the shore of Lake Issyk-
Kul’, the raccoon dog population reaches 20—25 animals рег 1000
hectares (Berens and Yanushevich, 1956 and others).

Habitat. In the Far East, preferred habitat of the raccoon dog is
light deciduous and mixed forests near streams with dense understory,
or the thick growths of shrubs, usually found on gentle slopes of moun-
tains, interrupted by waterfalls and creeks, with rock outcrops and clear
areas. In southern Ussuri territory, more than half of all encounters
with raccoon dogs, their burrows and excrements, were found in rare
oak-hornbeam forests with ground cover of rhododendron, honeysuckle
and other bushes. They avoid coniferous forests and are only encoun-
tered there along forest edges, riparian shrubs, or on cliffs along the sea
coast. In unforested regions, they are met with only in stands of reeds
along the shore of rivers and lakes. They usually live at an elevation
of up to 100—300 т above sea level, rarely up to 600—700 m (Bannikov,
1964).

In the central zone of the European part of the USSR, they prefer
deciduous and mixed forests of flood plains, and avoid dense coniferous
tracts and extensive open bogs and fields. At the northwest, the main
sites of the raccoon dog consist of southern taiga forest (30% of occur-
rences in summer and 49.5% in winter), shores of water bodies (26 and
20.8% respectively), bogs (12.7 and 10.2%), clearcuts and burns (11.3
and 7.1%); in this, mixed forests adjacent to floodplain sections of the
meadows and fields provide the principal sites both in winter and in
summer (Geller, 1959). In Novgorod district in flooded forests, the
number of raccoon dogs is 13; in grassy fir groves, 8; in pine woods,
1—3 animals рег 1000 hectares (Morozov, 1948). In Tatariya in forests,

ИЯ

Fig. 24. Reedgrass and sedge meadows in the lowland of Lake Khanka (southern
Primor’e). Biotope of raccoon dog. Photograph by A.G. Pankrat’ev.

raccoon dog encounter frequency is 33% (from 25 in summer to 40—
60% in winter and spring), in floodplains—30% (56% in summer,
10-19% in winter and spring), in ravines with bushes—11.5%, in
bogs—6.2%, in fields—14.4% (Yu. Popov, 1956). In Voronezh dis-
trict, they prefer to live in deciduous and mixed forests, especially on
floodplains where nearly half of all litters were noted; in small glades
in boggy depressions (more than 1/3 of the litters counted); in large
tracts of pine, they are confined to forest edges, in burns and clearcuts
(Obtemperanskii, 1953). In Gor’kovsk district, they prefer the flood-
plain deciduous forests with rich stands of grass (Kozlov, 1952). In
Byelorussia its favorite habitat is broad-leaved and mixed forest is-
lands, the copses among meadows and small bogs with dense
undergrowth of hazel, buckthorn and other bushes, and also dense
growths of alder thickets and oiser-beds along the banks of rivers and
lakes; it decidedly avoids dry pine woods (Dol’bik, 1952).

In Ukraine, it prefers the floodplains with growths of oiser and
high grass as well as the edges of bogs and flood-pools in the lower
reaches of large rivers. It establishes itself on river-terrace forests of
aspen and alder or in oak-groves, especially with dense understory. In
winter it does not abandon river floodplains (Korneev, 1954). In the
Donbass, besides floodplains, it willingly colonizes ravines overgrown

Fig. 25. Biotope of raccoon dog—Borzi river valley in its lower reaches, Amursk
district. Photograph by A.G. Pankrat’ev.

with shrubs, and in small shrubs in the steppes (Sakhno, 1948). In

_ Rostov district, it lives among shrubs, in ravines and depressions as

well as in growths of reeds (Rall’, Kritsksya, 1958). On the shore of
the Azov Sea, it is found in reed thickets and in growths of shrubs
(Kostyunchenko, 1950). Similar places are inhabited by this species
along the shore of the Caspian Sea and in the Volga delta. In the
northern Caucasus, the raccoon dog lives along the banks of rivers,
lakes and creeks, in ravines and gullies overgrown with bushes, in
isolated oak-groves among the steppes and in broad-leaved and mixed
montane forests. In the belt of montane coniferous forests, they are
encountered half as often as in mixed and broad-leaved forests
(Trushchalova, 1959). In Georgia, this species is confined to cut-over
forests and shrublands along river banks (Meladze, 1947); in Kirghizia,
they live along swampy shores of lakes, rivulets and in berry-bushes
and reeds (Berens, Yanushevich, 1956). The raccoon dog was intro-
duced into Siberia, but quickly died out; it inhabited forested floodplains
of rivers where nut-pine is found and shrubs are abundant; in summer,
they frequently. occupied open nut-pine woods in searching for food
(Nadeev, 1940).

Apparently, mixed and deciduous forests, alternating with small
open areas near water, represent the most favorable habitat of the

raccoon dog in its natural range, as well as in places where it has been
successfully acclimatized. In the majority of northern and central re-
gions of acclimatization, the raccoon dog prefers, in the snowy period
of the year, to live in open places (fields, meadows, marshes) where it
can move easily.

Food. According to the constituents of its food, the raccoon dog is
an omnivorous animal. The following morphological characteristics attest
to its omnivory: weak development of canines and carnassial teeth, flat
surface of molar teeth, and relatively long intestine (1.5—2 times longer
than that of other species in the family). The diversity of food objects
is very great. Animal food appeared in all stomachs and feces, while
plant food was, on the average, 64%.

Among animal foods, the most significant frequency of appearance
(in %) in all seasons and regions on the average are:

Insects 47 (maximum, to 93)
Mouse-like rodents 4685)
Amphibians 18-(65)
Birds 13—(23)
Fish 9-(24)
Reptiles 8(36)
Molluscs 747)
Carrion 613)
Insectivores 5—(14)

The absolute significance of one or another food is not only in terms
of frequency but also volume. In total, the role of mouse-like rodents
is considerably greater than that of insects. For example, the frequency
of occurrence of mouse-like rodents and insects in the food of the
raccoon dog of Voronezh preserve constitutes 60 and 46% respectively.
The volume of food (“food coefficient”—the product of the number of
occurrences multiplied by the volume in percentage, divided by 100)
for mouse-like rodents is equal to 58, and for insects—18 (Ivanova,
1962). In all regions inhabited by the raccoon dog, it may be confirmed
that mouse-like rodents occupy the first place in the nutrition of the
species. The significance of other food groups is very variable and
depending on the season and the conditions of the regions, insects ог
amphibians, birds or plant food constitute the food base. In one way or
other, all the above-mentioned food groups are always the main com-
ponent of raccoon dog food.

Insectivores Е tere ey ЗИВНЫНИ
| Mouse-like rodents | like rodents

Mouse: ike rodents

Reptiles
Amphibia

Fig. 26. Change in food of the raccoon dog by seasons (from Yu. Popov, 1956).

Mouse-like rodents in the food of the raccoon dog are mainly
represented by voles. Among the latter, those species living in damp
brushy undergrowth, banks of water bodies and coppices are more
frequent than other species; these are: reed voles, root voles, water
voles, common voles and others. In forest regions, the beasts rarely
feed on common redbacked voles. Mice are 5—10 times rarer than
voles; as an exception (in Astrakhan district) gerbils are of great im-
portance in the nutrition of raccoon dog.

Insects constituting the food of the raccoon dog are more frequently
represented by: dung beetles, cockchafers or water beetles and water
scavenger beetles; other groups are more rarely found: tiger and ground
beetles, carrion beetles, grasshoppers and others. Frogs are more com-
mon than the other amphibia among the food of raccoon dog. In some
instances, for example in Voronezh region, fire-bellied toads [Bombina]
were the usual food (Ivanova, 1962), while in the Ukraine, their usual
food was the spade-foot toad [Pelobates] (Korneev, 1954).

Among birds in the stomachs and feces of raccoon dog, the most
frequent are the remains of either ducks or passeriform birds; com-
monly passeriforms nesting in the area (buntings, pipits, warblers and
others) or else migrant species—thrushes, jays, etc. Where there are
colonies of nesting terns and gulls, their eggs and nestlings are common
food of raccoon dog. In the northern regions of raccoon dog acclima-
tization, grouse are usually its prey (Sorokin, 1956). In the Ussuri
territory, cases of destruction of pheasant nests were recorded.

The raccoon dog usually utilizes only large fish which are cast up
in spring after freezing; it rarely catches fish in passage during the
spawning migration. In Primor’e, it usually collects fish and inverte-

90
80
70
60
30
40
30
20
10
123456 123456 123456 123456
Spring Summer Autumn Winter

79 Fig. 27. Seasonal variation of food groups in the nutrition of raccoon dog (in
food-units) (from Ivanova, 1962).
1—Mouse-like rodents; 2—Carrion; 3—Birds; 4—Amphibia; 5— Insects;
6—Plant food.

80 Fig. 28. Raccoon dog searching for food on river ice. “Kedrovaya Pad’ ” preserve,
southern Primor’e. March 1964. Photograph by A.G. Pankrat’ev.

brates on the sea shore. It catches small fish from water bodies drying
out in summer. Snakes are the more frequent prey among reptilians.
Lizards are 2—3 times rarer. In the southern regions of their acclima-
tization, the raccoon dog willingly catches young tortoises and digs out
the clutches of tortoise eggs (Rukovskii, 1950). Both water molluscs—

х Ро —— ff Adult males
= 2000 ue —-—pp Adult females
1000 Be --- Juveniles

o-Vummuxuomigcow
Months

Fig. 29. Change in weight of the raccoon dog during the year (after Yu. Popov,
1956).

bivalves, pond snails and flat-coiled snails—and land snails are in-
cluded in the food of raccoon dogs.

Among insectivorous animals, the usual prey are shrews and hedge-
hogs, rarely moles and desmans. In the Far East, the Ussuri mole
[Mogera robusta] is the usual prey of raccoon dog, a fact which is
probably determined by the surface position of its feeding burrows.

Plant food is also highly variable. It may be the vegetative parts
of plants, bulbs, rhizomes, grains of agricultural crops (most frequent
are oats, millet, maize), nuts, fruits, berries, grapes, melons, watermel-
ons, pumpkins, tomatoes, etc. In Ussuri territory, the most important
are fruits of wild fruit trees, Amur cork trees, acorns and wild grapes.
Fruits of [cultivated] fruit trees and grapes have a great importance in
the feeding of the raccoon dogs acclimatized in the southern regions.
In the northern regions oats, berries (bilberry, cranberry, strawberry,
and others) and the vegetative parts of plants play a great role.

The fairly evident seasonal change in the food of the raccoon dog
is greatly determined by the different availability of these or other food
items during the year. Thus in late autumn and winter, mouse-like
rodents appear to be the main food of the raccoon dog, or in the north
it might be carrion and feces; since amphibians, reptiles, molluscs and
insects disappear. In the southern regions, the most important food
items are the grains of agricultural crops, fruits, and also those am-
phibia not undergoing hibernation (Korneev, 1954; Trushchalova, 1959
and others).

100

In spring, especially during snow melt, the number of mouse-like
rodents available to the raccoon dog sharply increase because of the
poor protection afforded by their burrows. During this period, amphib-
ians play also their greatest role. During the flight period of cockchafers
and the reproduction of dung beetles in the late spring, these insects are
also shown to be main foods. Vegetative parts of plants are very im-
portant. In summer the food is most diverse and, besides mouse-like
rodents, the relative role of which decreases, birds are of great impor-
tance (especially in the nesting period). Insects are important, and fish,
reptilians and molluscs are found more frequently than in other seasons.
By the end of summer, a greater role is taken by berries, grains, fruits
and vegetables.

In autumn, plant food is the most important item in comparison to
all other seasons. In connection with the increase in the number of
mouse-like rodents, their role increases, but the insects, amphibians,
reptiles and birds gradually disappear from the food ration (Table 1).

The importance of one or another food group differs remarkably
according to the year. Thus, in the food of raccoon dog, the frequency
of occurrence of the main food group—mouse-like rodents—may change
nearly two-fold (Table 2). The decrease in the role of mouse-like ro-
dents in the food (determined by the drop in their number) entails a
sharp increase in the significance of birds, amphibians and insectivores.
In other instances, the lack of mouse-like rodents is compensated by the
intensive eating of carrion (Novgorodsk and Kalininsk districts; Geller,
1959), feces (Tatariya; Yu. Popov, 1956), or fruits (Caucasus;
Trushchalova, 1959). Parallel to the decrease in the role of mouse-like
rodents in the food of the raccoon dog, the general significance of plant
food also decreases.

The food contributions of animals inhabiting different places also
vary somewhat. Thus, mouse-like rodents (common voles) are highly
important in the food of raccoon dog litters at the forest edges in
Voronezh preserve. Because of the low number of mouse-like rodents
within the forest, their importance in the food of litters living in the
center of the forest was very low; the deficiency of rodents here is
compensated by amphibians and insects (Table 3). The food of litters
living in floodplains was more diverse; the most important were water-
fowl (Ivanova, 1962). Differences nearly as sharp as these in the food
components of raccoon dogs inhabiting various biotopes were noted in
Tatariya (Yu. Popov, 1956), in Belorussia (Samusenko and Golodushko,
1961) and in other regions.

101

Table 1. Seasonal change in raccoon dog food in Primor’e
(E.P. Spangenberg) and in the northwestern European part of the USSR
(frequency, %; Geller, 1959)

REED rare Primor’e territory Northwestern USSR
umber Ob cate Spring and Autumn, Spring and Autumn,
summer, 26 summer, 130
Type of food 97 214

Animals
Insectivores — — 5.6 iS
Mouse-like rodents 18.6 15.4 1.7 48.3
Birds and their eggs 8.3 — 16.8 4.1
Lizards 7:2 — — —
Snakes 4.2 -—- Тгасе —
Turtle eggs 5:2 — a —
Amphibia 65.5 15.4 52.8 19.1
Fish 18.6 23.0 1.3 10.0
Insects 32.0 23.0 123 10.0
Molluscs 47.3 11.3 Trace 10.0
Carrion Dell — 6.5 15.8
Feces — — 57 1.6
Plants
Nuts and grains 11.3 11.3 — —
Grass 26.8 46.1 57.2 —
Oat — — 3.7 68.3
Rice 4.2 — — —
Polygonum, millet, сот 2.1 19.2 — —
Berries 2.6 —- 6.6 10.7
Grapes — 46.1 — --
Apples —— 62.2 — —
Cork tree — 61.5 os —-
Acorns — — a= 5.0

The sharp differences in the importance of one or another food in
different seasons in various years and in various habitats make it
difficult to establish rules governing the variability of food components
(Tables 1, 2, and 3a).

In northern and middle latitudes, mouse-like rodents occupy first
place among animal food; in the south and west—insects. As one
moves southward, the role of birds and molluscs increases. Naturally,
geographic variability of food includes replacement of some food types
by others within one food group (different species of mouse-like

102

Table 2. Change in importance of foods of raccoon dog in various years
(frequency of occurrence, %)

Region, author, Voronezhsk Tatariya Northwestern
year district (Yu. Popov, European
(Ivanova, 1956) part of USSR

1962) (Morozov, 1956;
Geller, 1959)

Type of food

1958 1959 1949 1950-1941 1947

Insectivores 5.4 15.0 No data 0.9 6.9
Mouse-like rodents 74.6 49.7 59.3 76.0 40.3 22.3
Birds 13.5 332 No data 4.3 10.2
Reptiles 2.2 322 No data — —
Amphibians 12.4 239) 14.1 6.5 11.9 29.8
Insects 48.6 43.9 56.0 53.0 29.8 7.6

Plants 50.8 40.8 40.3 28.6 94.7 34.0

rodents, insects, berries and so on in different regions). The extremely
scanty material for the feeding of raccoon dog in the Far East does not
permit any [comparative] analysis of changes in food components of
acclimatized animals [elsewhere] [the changes mentioned by Geller
(1959) are confined to annual and seasonal variability of feeding].

Home range. Data about the size of the home range are almost
absent. The distance between the inhabited burrows, which may serve
as an indirect indication of the size of the home range, is extremely
variable. The burrows may be separated from each other by some
kilometers or (as in Primor’e and on Askol’d Island) very close to each
other. Burrows were distributed in colonies consisting of 4—S (up to 10)
each in Rostovsk district where the foraging area of the raccoon dog
reaches 10 km? (Stepanov, 1939; Rall’ and Kritskaya, 1953).

In the northwestern regions of acclimatization, judging by 19 tracks,
the length of the daily travel of the raccoon dog is equal to 3—6 km in
winter, in early spring 15—20 km, in late spring 6-8 km, in summer 6—
10 km, in early autumn 8—12 km, and in late autumn 2—3 km. The area
within which these travels take place, was equal to | to 12 km? (Geller,
1959). In Таапуа, the length of the daily travel of the animals in
search of food in early spring reaches a maximum of 10—14 km; in
summer, when food is abundant, this travel is less and the radius of
activity of seven raccoon dogs consisted of 600-800 т. In autumn,
when fat is accumulating, the length of the foraging circuit may reach
8 km, but usually is 4-6 km. In winter, during periods of thaw, they

103

Table 3. Foods of raccoon dog litters in different habitats in Voronezh
Preserve (frequency of occurrence, %)

Biotope Floodplain Forest Forest edge
Type of food (burrow No. 1) (burrow No. 2) (burrow No. 3)
Mouse-like rodents 25.7 5.0 56.0
Birds 48.6 — 20.0
ВерШез 8.6 10.0 8.0
Amphibians 31.4 70.0 16.0
Insects 62.8 100.0 52.0
Plants 25.7 20.0 56.0

do not go further than 100-150 m away from shelter (Yu. Popov,
1956).

Burrows and shelters. Raccoon dogs use several types of shelters:
beds and lairs in open areas, and burrows, both temporary and perma-
nent ones (breeding and wintering dens).

In the Ussuri territory, summer beds are generally places on dry
slopes under a protective rock, boulders or in brushy undergrowth. In
the Khankaisk lowland, beds/nests found in September—October were
between hummocks on a thick (4.5—6 cm) layer of accumulated cuttings
of dry sedge, reed and other grassy vegetation. At the bottom of the
nest, scraps of wool were found. The internal diameter of such nests is
78—100 cm, the depth is 32—36 cm. Around the nest the reeds usually
form an obstruction by arching above the nest. In the Ussuri territory,
beds are found in the depressions between roots of old tress and in the
spacious hollows of fallen lime trees [Tilia].

Based on investigation of 49 burrows in the Ussuri territory, the
majority of the holes are located 500—800 т away from the banks of
water bodies, most frequently (41%) at eastern points of the compass.
Slopes on which burrows were found were covered with forests (88%)
or shrubs (12%). 78% of the burrows were found under piles of stones,
12% in loose soil, 8% in rock niches, and 2% under tree-roots (Bannikov
and Sergeev, 1939).

The horizontal diameter of the passage of burrows beneath stones
is often equal to 24—29 cm (21 to 31 cm), vertical diameter 20-24 cm
(20 to 34 cm). In the burrow there are usually 1—2 entrance holes, in
some instances 3—5. The longest passage to the nesting cavity is 2 m,
more frequently 1.5 m, 1.е., the burrows of this type are comparatively
short and simply constructed. The nesting cavity has a flat bottom and
a dome-shaped roof; its dimensions are 50—70 x 30—45 cm.

104

059 сур вер ON

S = 67C
vl = ae
Гб Lae a
vv BSE GE
ak OSI 18

sajdures $16 `(6961

‘вло[еЧ2Ч$111) snsvoned WoyWON

sajdwes $85 (1961

sajdues 90] “(OS6l
‘eAeysyiy) soddajs 1$ЧэАиеи|

‘IPJSAOYNY) JMISIP ysueyFeNsSY

ges
Cal
8`0
v'L
VIC
BL
691
OLE
8°99
СЗУ

sojdures

00S “(PS6l ‘лээшоу) ouTeIy/

sojdwies 777

(IS61 ‘шрозоЯ) эллэзэзА IP{SIO

CUP 6°89
Gel 61
85 £0
9°0 81
Sat iG
a [0
0' 55 Сс
Li Sv
SOV 9`УР
709 v'97

‘(ZS61) WysuesoduisiqQ

sojduies 9¢9 `(596Т) елоцел]

0091

JOSIP
YSYZOUOIO A,

ViVi — СЕ LC

al = 8°81

8 Е ==
УТ Sl Oe
6 69 Oe
61 Sal [`0
6L то 0'SI
8°С1 ge ese
OCS 2105 Гус
УЕ 608 Е

sojdwies 7697

(9661 ‘Aodog "nA) eAuEye

USSN IY}? JO suoises yUIAIIp ш Sop иоозэвл JO иоцызпи

sojduies

sojduies 0$ “(1961 ‘охЧзпрооо
pue oyuosnues) eissniojahg
СЕТ “(0961 ‘эиезтаа) BlUeNYyWT

L’v6

€6°°0

DiC
РС
950
ЕР
te)
8°67
Les

sojdwies

9901 ‘`([96Т ‘AOZOIOW)) S}OLSIP

yspeisulus] рие ySpo1osaon

[УС ие
OE SOJOAIIOOSU]
С uowe)
50 SOSNI[OW|
01 sojndoy
vl Ч
76 зрия
6'61 sueiqrydwy
602 $12954]
8'05 $зиэрол эу|-э5п0и

9по18 poo,

sojdures $816

"(9661 ‘\14010$) JOLNSIP узиииех

dy) Ul $91048 рооу шеш ay) Jo (%) Аэиэпбэлд “wg э14в1,

€8

105

Fig. 30. Burrows of raccoon dog. Scheme of location and cross section (after
Bannikov and Sergeev, 1939).

Most often (36 burrows out of 38) there is only one entrance to the
cavity; the side burrows join together in front of the nesting cavity in
one principal passage. The bottom of the cavity is usually lined with
different dry plant remains, leaves of hazel, oak, maple, lespedeza,
stems and leaves of grasses and herbs.

Near the burrow, usually not farther than 10—15 m, and in the
burrows with litters, toilets in the form of pits are located, sometimes
directly near the entrance, filled with excrements. Near the burrow
under piles of stones occur latrines in the form of numerous heaps of
excrements on flat rocks. In such cases, the latrine is frequently located
20—30 т, or farther, from the burrow.

Burrows excavated in soft ground in Ussuri territory were situated
along the slopes of river valleys, in places poor in rock outcrops. They
are quite simple, having 1-3 exits averaging 21 x 24 cm in diameter
(18—22 x 25—29 cm). Fan-shaped scattered soil and gravel are found
in front of the entrance to the hole and lower along the slope. The
burrow passage usually passes downward directly into the ground without
any turns at an angle of 20—25°; its length reaches 6 т (most frequently
3 m) and ends with a nesting chamber which does not essentially differ
from that described above in burrows under stones, but in the holes dug
in soft ground, the layer of leaf and grass bedding is less. In some
burrows a blind side hole proceeds from the nesting chamber which
sometimes rises slightly upwards, in which the concealed animal is
found when the burrow is dug up. Burrows in rock niches are only
accessible for external inspection. There is one entrance to such bur-
rows; wide fissures (more than 50 cm) and those opening upwards are
not occupied by raccoon dogs (a burrow located under the roots of an
old lime tree had 3 entrance openings under the tree).

Entrance\No. 4

106

Chamber*
of the burrow

Bifurcation No. 1
-Light-gray loam д ;
/ Loam with j { sand

0 ais mg? 93

Fig. 31. Scheme of construction of raccoon dog burrow (from Yu. Popov, 1956).
*In Russian original kotel, which means “‘caldron, boiler, pot’—Sci. Ed.

Shelters of the raccoon dog in places of their acclimatization are
similar to those described. In Tatariya and Rostovsk district, summer
beds, in the form of small holes without bedding are situated on dry
places at the base of slanting shrubs growing in depressions, and the
shores of marshes and lakes (Rall’ and Kritskaya, 1953; Yu. Popov,
1956). The raccoon dog sometimes constructs a lair in tall, dense sedge
thickets in the form of a plaza (70 x 100 cm) on small piles of mowed
sedge or other grass remaining from the previous year. In Tatariya,
winter lairs on the frozen swamps in dense growths of reed and cattail
(Yu. Popov, 1956) resemble lairs in the Khankaisk lowland. Raccoon
dogs accumulate the dry sedge to serve as bedding, while the surround-
ing high grasses bend under the weight of the snow forming an arch
above the lair. Below it the animals construct a whole labyrinth of
hidden passages, ending in an outlet on the surface of the snow cover,
sometimes 15—20 cm away from the lair. Such lairs constituted in
Tatariya rhore than 37% of all aboveground shelters.

In many regions of acclimatization, typical lairs are under hay-
stacks and shocks. Such dens are usually also used in winter or for
whelping; they are situated under tangled fallen trees, under stones, in
old sheds, in trenches, in dug-outs, and in entrenchments remaining
after the war. In such dens, there is bedding of dry grasses and leaves.
Shelters in the hollows of fallen trees or in hollows located at ground
level are quite frequent. Thus, in the northwestern European part of
USSR, 17% of winter dens of raccoon dog were found under the roots
of trees, 13%—in dug-outs, 10%—in entrenchments, 3%—in hollows
and stacks (Geller, 1959). In Tatariya, 5.9% of the lairs were found in
hollows (burrows not included)—4.8% under haystacks and 5.3%—in
heaps of dry branches (Yu. Popov, 1956).

Entrance
No. 1

11.6

107

The significance of burrows as shelters for the raccoon dog in
places of acclimatization may be highly variable even in neighboring
regions. Thus, in Voronezhsk district, about 75% of the litters of the
raccoon dog were found in burrows (ОМетрегапзКи, 1952), in Voronezh
Preserve—more than 86% (Ivanova, 1962). In the latter, old burrows
of badgers and red foxes are especially numerous, and the ground is
sandy. In the northwestern regions, about 50% of the raccoon dogs
occupy burrows (Geller, 1959); in the Ukraine, they rarely utilize
burrows (Korneev, 1954), but in Rostovsk district they live, as a rule,
in burrows (Rall’ and Kritskaya, 1953).

The raccoon dog usually hurries to occupy the old burrows of
badgers and red foxes, and digs itself a new burrow only when there
are no free ones. In Tatariya, 77.8% of the burrows occupied by the
raccoon dog were old ones of badgers (38.6%) or of red foxes (39.2%)
(Yu. Popov, 1956). The samie phenomenon was observed in other re-
gions. In using the burrows of badgers and foxes, the raccoon dog, as
a rule, does not dig additional side burrows and does not construct a
new nesting chamber, but only brings in dry bedding, throwing away
the rotted. On occupying the complicated holes of badgers, the raccoon
dog uses 2—3 side burrows and the others are neglected.

Many cases of wintering of raccoon dog and badger in one hole
have been recorded. This is possible, because the badger enters into
hibernation at least 2 weeks earlier than the raccoon dog and leaves the
burrows 2 weeks after the raccoon dog. In the event that the raccoon
dog remains longer in the burrow, the badger drives it away and not
rarely bites it to death (Korneev, 1954; Geller, 1959; Ivanova, 1962
and others). Exceptionally, cases of cohabitation of litters of badger
and raccoon dog in one burrow are known (Tikhvinskii, 1938; Yu.
Popov, 1956 and others).

In those cases, when the raccoon dog itself digs, it constructs
temporary as well as permanent burrows. Temporary burrows are simple
in construction, small and short (length to 1 m), always with a single
entrance hole. The animals dig them in soft ground, often under bushes
or young pine trees, usually on precipitous slopes, along river banks,
at forest edges and in fields. Permanent littering and wintering burrows
are more frequently dug on the slopes of forest ravines, in floodplains
of rivers and at the borders of bogs, not far from forest edges or
openings in the forest and often on elevated parts of watersheds. In this
case, preference is shown for deciduous and mixed forests of various

108

species, with a rich forest understory. The nature of the ground is not,
apparently, of decisive importance, but sandy soils are preferred.

Proper burrows in Voronezh Preserve had one entrance hole in
57.1% of cases, two in 25% and more than 2 entrance holes in only
18% (Ivanova, 1959). This relationship of burrows with various num-
bers of passages is also found in other regions. The general length of
the burrow ranges from 1.5 to 5 m, often 2.5—-3 т. A chamber 40-50
cm in diameter is situated at a depth of 60—90 cm (usually 70—75 cm).
Width of the entrance hole exceeds its height, its dimensions being
24x29 cm (from 19 to 41 cm). One passage usually leads to the
chamber; as an exception, blind side holes occur, proceeding 60—80 cm
away from the chamber. The floor of the chamber is lined with a small
layer of bedding made of dry grass and leaves.

At the burrow, soil thrown out in a fan shape over 1.5—1.8 т
occurs. The dug-out soil and the area around the burrow are trampled
down by the animals, and evident traces of their thin claws are usually
seen on the walls of the burrow. 5—10 m, but sometimes 2—3 m, from
the burrow is located a latrine with pits. The trails of the animals are
seen on the discarded soil but these paths are narrow (not exceeding 20
cm), and this distinguishes the holes of raccoon dog from those of
foxes, the paths of which are broad (25—35 cm) at the burrow, and the
excavated soil coming from the hole is rounded in shape. The burrow
of the raccoon dog differs from those of the badger in that the soil
thrown from the hole of the latter is elongated, and the trails are not
usually seen, since all the excavated soil is trampled down by the feet
of the animals (Yu. Popov, 1956; Ivanova, 1959, 1962 and others).

The raccoon dog, apparently, changes the shelters seasonally. Thus,
in Tatariya, the beasts overwinter in burrows, dens or winter beds.
With increased activity in March, they often start to use temporary
burrows or beds, but at the end of April, before whelping, they again
occupy permanent holes, throwing out the old bedding from them and
bringing in new. By the middle of June, when the pups are about one
month old, the animals leave the littering shelters and use temporary
summer shelters, only rarely visiting the burrows. In autumn, visits to
the permanent holes become frequent, and in the middle of October, the
animals clean them a second time and change the bedding; at the end
of October, they finally settle in their winter shelters (Yu. Popov,
1956).

Daily activity and behavior. The raccoon dog is a nocturnal and
crepuscular animal. However, in the short nights of summer, activity

109

occurs during sunny periods in the morning and evening hours. During
the day, activity of the animals is curtailed and they hide in shelters or
spend the time bedded down. During the period of rut, pregnancy and
the initiation of nursing the young, from March to May, the raccoon
dogs are very cautious and are active almost exclusively in the dark
périod of the 24 hours. In summer, when the pups begin to feed inde-
pendently, they frequently are encountered during the daytime. In autumn,
in September, regardless of increased feeding, raccoon dogs rarely
come out in the daytime and usually leave their shelters only at twi-
light. In winter, the animals are active at twilight and during the night.

Upon leaving the burrow the raccoon dog in search of food follows
the shore of the nearest water body. It frequently goes into water and
wanders along the shallows.

On the sea coast, along shores of lakes and rivers, at dawn and in
the evening, it is almost always possible to see feeding animals. In the
forest the raccoon dog moves slowly, looking under every bush, pile
and stump. Only when crossing openings does it move at a quick trot.
The raccoon dog is cautious but in case of danger it does not run away,
but conceals itself, camouflaging itself very skillfully. If they are over-
taken, some animals (in the Far East) pretend to be dead, but on the
first opportunity, they run away and conceal themselves nearby. The
entire litter usually grows fat together at the end of summer. It is
possible to observe two or three adult animals, feeding in immediate
proximity to one another.

Having comparatively short legs, the raccoon dog, in the presence
of a snow cover 10—12 cm deep, already stops “dragging”, and searches
for food with difficulty. In a snow depth of more than 20 cm deep or
deeper the animals move with great difficulty, “swimming” in the snow.
The weight load for the track is relatively great: in December it is 75—
80 g/cm’, in March 33—37 g/cm? (Yu. Popov, 1956). The sense of smell
is very sharp, and hearing and vision are more weakly developed than
in closely-related species.

Hibernation, winter sleep. True winter torpor, i.e. deep uninter-
rupted sleep, associated with a sharp fall in metabolic level and body
temperature, is not observed in the raccoon dog. However, the general
metabolism may decrease by about 25% (Sokolov, 1949).

In warm winters in the Ussuri territory, as well as in places of
acclimatization (Ukraine; Korneev, 1954), the beasts do not go into
hibernation during the course of the entire winter and only during
snowstorms they do not go out of their shelters for some days. Usually

110

in December, when the snow cover reaches 15—20 cm, the activity of
the raccoon dogs sharply decreases and they leave their shelter for a
short time only on warm, quiet days and they do not go farther from
the burrow than 150—200 т. From February to the beginning of March,
the activity of the animal again sharply increases, both from food
deficiency as well as the beginning of rut, and this period leads to an
increase in the movements of the raccoon dog. During spring snowfalls
(March—April) the animals again may go into shelter for some days.

At the beginning of the winter period, the raccoon dog puts on an
average of 18-23% of subcutaneous and 3—5% of internal fat (of total
weight). Animals putting on less than this amount (though often gain-
ing) as a rule remain wandering and rarely survive the winter. During
the first years of acclimatization, when animals were not yet suffi-
ciently adapted to catching the local food, as a result of this they
exhibited weak fattening in autumn, and frequently did not hibernate
(Obtemperanskii, 1958 and others). Low temperature exerts little influ-
ence on the activity of the raccoon dog. Instances are known of animals
being encountered at —20 to —25°C.

Raccoon dogs kept in artificial burrows in hutches fell into hiber-
nation in December, but aroused and fed during a time of warming; in
the period of rut (in mid-February) they wandered about, but then at
several times hibernated again (Obtemperanskii, 1958). Confirmation
of partial winter activity of the raccoon dog is attested by their being
caught by hunters. Thus, in Tatariya, about 80% of the animals are
caught from November to January and 20% from the Ist of January to
the 15th of February (Yu. Popov, 1956). In the northwest of the coun-
try, about 37% of the animals are caught in December, in January—28%
and in February—18% (Geller, 1959).

Seasonal migrations and transgressions. The raccoon dog is a
sedentary animal, and regular seasonal migrations are not known for
this species.

In the Ussuri territory, hunters have reported migrations of raccoon
dogs in unfavorable years. In the Lake Khanka depression, massive
migration of the animals to China seemed to have been observed in the
autumn of 1929 when, within a few days, up to one hundred animals
were seen moving in one direction. Such migrations are doubtful; prob-
ably what occurs is the migration in the winter of a small number of
animals living during the summer in reeds from flooded depressions to
dry places.

111

Fig. 32. А young raccoon dog, just appearing after winter sleep. “Kedrovaya Pad”
preserve, southern Primor’e. 17 February 1965. Photograph by A.G. Pankrat’ev.

After releases in new regions, there were migrations of individual
animals for 20-80 km from the place where they were originally intro-
duced (Rall’ and Kritskaya, 1953; Trushchalova, 1959 and others).
There is the possibility of more distant migrations, since intrusions of
the animals onto the tundra have been recorded (Skrobov, 1958). An
emigration of about 600 km was recorded in Ukraine (Korneev, 1954).
The greatest migrations of the raccoon dog are recorded in floodplains;
during the final spring floodwaters, females with litters move up to
divides.

Reproduction. Rut is observed from the beginning of February to
the end of April depending upon the region and climatic conditions.
Later spring may delay the beginning of rut for 2—3 weeks. The return
of cold weather, and especially snowfalls, usually interrupt rut. In the
Ussuri territory, rut takes place on the average in the middle of March,
but it often is delayed until the end of March or beginning of April. In
the Ukraine, in Voronezhsk district and Tatariya, rut occurs in the
middle of February (Korneev, 1954; Yu. Popov, 1956; Obtemperanskii,

1958); in the northwest, in the beginning or middle of March (Geller,
1959 and others). Rut continues for 2—3 weeks, but in captivity, it is
usually not less than 3 weeks, and often 26—28 days, and during food
shortage, up to 40 days (Pavlinskii, 1937).

Raccoon dogs are monogamous animals and pairs are formed even
in autumn, usually in October or November. Fights among males occur
occasionally at the time of rut, are brief and not violent, and are
accompanied by yelping and hollow growling. In captivity, polygamy
is possible when one male is in contact with 4—5 females. Copulation
occurs most frequently during the night or early in the morning, usually
in quiet weather. Coitus lasts for 6-9 minutes on the average (from 2
to 26 minutes). Estrus in females continues from several hours to 6
days and during this period they mate up to 5 times, usually 2—3. After
20—24 days, estrus is repeated even in pregnant females (Petryaev and
Khatkevitch, 1931; Pavlinskii, 1937).

Pregnancy lasts 61—70 days (often 59 days [sic]*). Young are born
in April, but mostly in May. Cases are known when the main mass of
births is delayed until June, and in a few cases newborn have been
found as late as September.

In the Far East, the litter mostly consists of 6-7 young, but their
number may reach 15—16 (Firsov, 1929; Przheval’skii, 1870**). The
size of the litter in the regions of acclimatization fluctuates greatly, in
connection, probably, with fluctuation of conditions in various years.
Thus, in Voronezhsk district (Obtemperanskii, 1958) average litter size
was 4.9 pups (from 4.3 to 5.75 in different years); in the northwestern
districts in 1938—1939—5.2; in 1947—5.9; in 1946—1949 average, 6.3
(Morozov, 1953; Geller, 1959). In Tatariya (Yu. Popov, 1956) the
average litter size was 7.3 pups; in the Ukraine (Korneev, 1954)—7.9
(to 16 pups); in Lithuania—from 4 to 13, average, 9.5 (Prusaite, 1961).
The average occurring in captivity is 7.1 pups (Petryaev and Khatkevitch,
1931). Females giving birth to pups for the first time have smaller
litters than females of older age, and their litters are 1 or 2 pups fewer.
The ratio of sexes at birth is nearly 1 : 1; but males are somewhat more
frequent. Thus, in Tatariya (Yu. Popov, 1956) litters contained 50.9%
males and 49.1% females; in the northwest (Geller, 1959}—51.7% and
48.3% respectively; and in Voronezhsk district (Obtemperanskii,
1958)—58.7% and 41.3%. In State animal farms (Povlinskii, 1939)—
52% males and 48% females.

*Russian original inconsistent—Sci. Ed.
**In Russian original, 1947 (not in Lit. Cit.—Sci. Ed.

E43)

Barrenness of females in different regions of acclimatization con-
stitutes about 12% (Yu. Popov, 1956; Geller, 1959 and others). In
captivity, after one mounting [by male] 30.9% of the females remain
non-pregnant and after three mountings, practically no females are
barren (Starkov, 1940). The male takes an active part in raising the
young. In Tatariya the average number of pups in the litter decreases
from 7.3 in May to 5.9 in June due to mortality (Yu. Popov, 1956).
Juvenile mortality increases in June when the pups leave the shelter (at
an age of 25—30 days), and reaches its maximum in July and August
when the growing pups separate from the litter. The litter finally breaks
up at the end of August-September, and in October juveniles, reaching
the size of adults, unite in pairs. During this period about 35% of the
young have died, and each pair of adults during this time produces 4
juveniles (Morozov, 1953). By September and October, about 50% of
the young have died (Geller, 1959).

Growth, development, and molt. Young are born blind, covered
with short, dense, soft wool, without guard hairs, of dark-slaty, nearly
black color. Weight of newborn, 60—110 g. Males weigh 5—10% more
than females. Eyes open on the 9th—10th day. On the 14th—16th days
teeth erupt—beginning with the upper canines, then the incisors and
lower canines. Approximately from the age of 10 days, guard hairs
begin to grow on the hips and shoulders, and then near the ears and on
the cheeks. The dark color of the pup gradually becomes gray, starting
with the abdomen, 2 weeks after birth. Guard hairs start to grow
especially quickly at the age of 1.5—2 months when the black or black-
ish-brown color prevailing in the color of pups quickly changes to gray.
The black tones remain only around the eyes. On all the interorbital
area, from the forehead to the nose, the brown color of the underfur
also changes into light gray. By this time, as a result of lengthening of
hairs on the cheeks, “side-whiskers” begin to form. In 3-month-old
pups, the guard hairs are shorter than those of adults only on the chest,
chin, tail and distal parts of the extremities.

Lactation lasts for 45—60 days, but from 3 weeks or one month of
age, the pups start eating frogs, insects and other food brought to them
by the parents. At the age of | month, the pups weigh 550—650 g; at
2 months, 1100—1300 2; 3 months, 2500—2900 2; and at 4 months —
about 4 kg. At an age of 4.5—-5 months, young attain the weight and size
of adults and are almost not different from them in color (Pavlinskii,
1937; Korneev, 1954; Yu. Popov, 1956; Geller, 1959 and others).

The lowest weight of adults is attained in March (about 3 kg). By

114

August—beginning of September, males reach their greatest weight—
6.5—7 kg (certain individuals, 9-10 kg). Females deposit fat reserves
more slowly and maximum weight is usually attained one or one and
а half months later than males, 1.е. by September—October. Juveniles
attain greatest weight another month later than females—in October—
November. In autumn, fat constitutes 30-35% of live weight of the
animals and forms a subcutaneous layer up to 1.5 cm (Yu. Popov,
1956; Geller, 1959).

Sexual maturity is attained at an age of 8—10 months. The duration
of life is not known. In nature, individuals older than 6—7 years are
encountered. In captivity, animals are known to have lived to 11 years.

The age composition of populations is insufficiently studied. In the
northwestern regions, juveniles constituted 53.8% of the population in
the hunting season and animals of older ages—46.2% (Geller, 1959).
Among animals caught in Tatariya in the hunting season, juveniles
constitute 57.2% of the population, adults—42.8% (among them, 19%
are older than 1 year, 11.3% older than 2 years, 6.7% older than 3
years, 3.9% older than 4 years, 1.9% older than 5 years). In Lithuania
in 1957—1959, one-year-old animals constituted 67.5% of the popula-
tion (Prusaite, 1961).

Molt occurs only once a year—in spring. Shedding of the underfur
starts in February—March depending upon the region, coming of spring,
and condition of the animals. Shedding of underfur starts on the neck
and withers; then molt extends to the shoulders, back, sides and the
posterior part of the body. By the middle of May—June (and July in the
northern regions), guard hair almost alone remains on the skin, and the
fur acquires a dark color because of the proximity of the tips of the
guard hairs, which have black tips. From June onward, guard hairs are
gradually replaced and the underwool starts growing. Renewal of the
latter takes place slowly compared with guard hair, and therefore the
pelage attains normal development from the last days of October, to
November, and even December. The underfur (new) per 1 cm? of the
rump is in August—2072, in October—6264, and in December—9624
(Trushchalova, 1959). In old males, molt takes place more quickly than
in females and juveniles (Korneev, 1954; Morozova, 1955; Geller,
1959 and others).

Enemies, diseases, parasites, mortality, competitors and popula-
tion dynamics. Among the enemies of the raccoon dog, the most important
is the wolf. It destroys raccoon dogs early in spring—in February—
March, and also in summer. However attacks on raccoon dogs by

—

115

wolves have been noted in late autumn also. Stray dogs destroy mainly
the growing pups. Apparently, during the period of litter dissolution,
red foxes also destroy pups. Instances are known of foxes biting rac-
coon dogs to death (see below).

In Tatariya, out of the 54 recorded cases of death of raccoon dogs
by predators, 55.6% of the animals were killed by wolves, 27.8%—by
stray dogs, 11.1%—by red foxes, 3.7%—by the golden eagle and
1.8%—y the eagle-owl [Bubo bubo] (Yu. Popov, 1956). In the north-
western regions, out of the 186 recorded cases of death of raccoon dogs
by predators, 64% were attributed to the wolf, 14.3%—+to fox, 12.8%—
to stray dogs, 6.3%—to lynx and 2.6%—to birds of prey (Geller,
1959). Lynx, because of its low numbers, causes little damage to the
raccoon dog, as observed in the majority of regions; still more rarely
do brown bears attack raccoon dogs. Among birds of prey involved in
attacks, usually on pups, are golden eagle, white-tailed eagle [Haliaetus
albicilla|, goshawk and eagle-owl.

Among the raccoon dogs’ competitors, fox and badger are the most
important, since foxes play a role with raccoon dogs in feeding on
mouse-like rodents, whereas the badger generally shares with them
feeding on insects and plants. However in the summer period, thanks
to the abundance of food, the various habitats and different methods of
hunting, there is no sharp competition among these species of predators.
This competition becomes somewhat sharper in early spring (February,
March), in the most difficult period of the year. Competition with foxes
and badgers for burrows is practically absent, for the raccoon dog is
entirely unassuming in its struggle for shelter (see above). However,
individual cases have been recorded, when badger or fox killed the
raccoon dog which had occupied its burrow.

The parasitic fauna of raccoon dogs was investigated in the Far
East (Shpringol’ts-Shmidt, 1935; Petrov, 1941), in State animal farms
(Gusev, 1951; Savinov, 1953 and others), in Tatariya (Yu. Popov,
1956), in the Ukraine (Korneev, 1954), in the northwestern regions
(Antipin, 1946; Morozov, 1953; Savinov, 1953; Geller, 1959), in the
northern Caucasus (Trushchalova, 1959), in the lower Volga region
(Sviridov, 1952), and in Lithuania (Prusaite, 1961).

In ail, 32 species of parasitic worms are known from the raccoon
dog: 8 species of trematodes (Opisthorchis felineus, Clonorchis sinensis,
Pseudamphistomum truncatum, Euparyphium melis, Metagonimus
yokogawai, Plagiorchis massino, Alaria alata, Paragonimus
westermani), 7 species of cestodes (Diphyllobothrium mansoni,

116

Taenia hydatigena, T. polyacantha, Multiceps multiceps, M. serialis,
Diphylidium caninum, Mesocestoides lineatus) and 17 species of nema-
todes (Ascaris columnaris, Toxascaris leonina, Т[охосага] canis,
СарШата yamaguti, С. plica, С. putori, Thominx aerophilus,
Ancyclostoma caninum, Uncinaria stenocephala, Molineus patans,
Crenosoma vulpis, Dioctophyma renale, Physaloptera sibirica, Thelazia
callipaeda, Strongyloides _erschowi, ТисйтеЦа _ spiralis,
Macracanthorhynchus catulinum*).

In the Far East, 18 species of parasitic worms were found in
raccoon dogs, and 12 species were found in regions of acclimatization.
Such species as Clonorchis sinensis, Metagonimus yokogawai,
Paragonimus westermani were not found in acclimatized animals due
to the absence of their intermediate hosts in Europe—the Eastern mol-
luscs. Some species: Opisthorchis felineus, Plagiorchis massino, Taenia
hydatigena, T. polyacantha, Multiceps multiceps, M. serialis, Ascaris
columnaris, Ancyclostoma caninum were acquired by raccoon dogs
during the process of acclimatization.

In the northwestern regions there has been noted a high degree of
infection (up to 96% and 10—20 parasites per 1 cm? of small intestine)
with the trematode Alaria alata; somewhat less (33.7%)—with the
nematode Uncinaria stenocephala (Savinov, 1953; Geller, 1959). In
Tatariya, a great number of animals were found to be infected by
Alaria alata, but the total number of infected animals was less (76.2%)
than in the northwest (Yu. Popov, 1956), in Lithuania, a lesser number
of animals (54%) was infected by this parasite (Prusaite, 1961). In the
Ukraine, 3 species of parasitic worms were found, including Alaria
alata in 33.3% of animals. Infection with Alaria takes place through
frogs infected with the metacercaria** of Alaria. In the Caucasus
(Trushchalova, 1959) a considerable number (27.9%) of raccoon dogs
are infected with the ascarid Ascaris columnaris. 9.3% of raccoon dogs
of Lithuania (Prusaite, 1961) are infected with the larvae of Trichinella
spiralis. However, the pathogenic significance of parasitic worms to
raccoon dogs is little studied. In Kievsk district in 1952, there was
massive mortality among raccoon dogs due to bronchopneumonia of a
trematode nature (Korneev, 1954).

Six species of fleas have been found on raccoon dogs: Chaetopsylla
trichosa, C. globiceps, Paraceras melis, Ctenocephalides canis,

*an acanthocephalan; misspellings in Russian original have been corrected—
Sci; Ed.
** Actually, mesocercaria—Sci. Ed.

117

C. felis, Pulex irritans, as well as chewing lice, Mallophaga; 5 species
of ticks: Dermacentor pictus, Ixodes ricinus, I. persulcatus, I.
crenulatus and the causative agent of mange—Acarus siro*. The latter
is fairly widespread in the Ukraine, Voronezhsk district, Lithuania and
a series of other regions, causing massive outbreaks of mange. Raccoon
dogs are strongly infected with the biting louse Trichodectes canis; in
the State animal farms of the Far East, this parasite afflicts up to 90%
of animals (Shpringol’ts-Shmidt, 1935) causing a marked exhaustion of
the animals.

Diseases of the raccoon dog are poorly studied. Cases of rabies
were recorded in the lower Volga (Isakov, 1949), Voronezhsk district
(Obtemperanskii, 1958), Lithuania (Prusaite, 1961) and in other
regions. In the Ukraine (Sakhno, 1948; Korneev, 1954) and in Tatariya
(Tikhvinskii, 1938) cases of massive epizootics of piroplasmosis were
recorded. In the northern Caucasus, epizootic carnivore distemper was
noted among raccoon dogs (Trushchalova, 1959); a known case of a
raccoon dog ill with leptospirosis (N.N. Rukovskii); in State animal
farms, carnivore plague, paratyphoid, anthrax, and tuberculosis sick-
ness (Burov, Buzinov, Bannoviskii, Lyubashchenko and others, 1939).

Epizootics may be an important factor in reducing the number of
raccoon dogs. “An almost complete destruction of the whole stock” was
observed as a result of an epizootic of piroplasmosis in Kievsk and
Chernigovsk districts (Korneev, 1954) and in Tatariya (Tikhvinski,
1937). A great number of raccoon dogs are lost to mange, which, |
however, does not lead to a massive die-off of the animals, as is known
for foxes (Prusaite, 1961 and others). The number of raccoon dogs may
be sharply reduced as a result of mortality from rabies (Isakov, 1949).

Among the abiotic factors which cause a sharp decrease in the
number of raccoon dogs, the spring flooding which coincides with the
period of reproduction is important. At that time, young die in the dens;
and, apparently, a significant number of pregnant females are killed
(Korneev, 1954 and others). In the northern regions, a cause of reduc-
tion in numbers may be spring snowfall, involving lack of food, death
due to predators, increase in the number of non-pregnant females and
lowering of fertility (Yu. Popov, 1956; Obtemperanskii, 1958; Geller,
1959 and others). There are no concrete data about the magnitude of
mortality due to different causes. In the flooded areas of the Dnepr,
Volga and other large rivers, in some years the greater part of popu-
lation is killed. However, in favorable years, due to the high fertility

*Mange is also caused by Sarcoptes—Sci. Ed.

118

(see above) the population doubles by autumn, and the number of
raccoon dog is quickly restored.

The number of raccoon dog is related to food availability. Thus, in
Voronezhsk district, 3—4 fold fluctuations in the number of this carni-
vore were observed. The fall and rise in the numbers is related to
changes in the numbers of the principal food—mouse-like rodents —
in the preceding year and has a 34-year cycle (Ivanova, 1962).

Field characteristics. A stocky animal of medium size, on-short
legs, with small head and pointed muzzle; ears almost hidden in thick
fur and typical “side-whiskers” on the cheeks.

When in danger, conceals itself, closely presses itself to the ground
and, owing to its brown color, merges with the surrounding soil back-
ground or forest bedding. In case of direct close approach of a human,
it usually closes its eyes and lies completely still, even when touched.
During twilight and night, it is more courageous and sometimes tries to
defend itself. It swims well, willingly enters water and can swim across
wide rivers and lakes.

In the places inhabited by raccoon dogs, holes dug in search of
insects may be found. These holes are 5-15 cm in depth and are

| и
Ф Fi 4
® Е, vine

Fig. 33. Prints of front and hind paws of raccoon dog in silt. On the right, scheme
of the usual position of tracks during slow movement. Volga delta, 24 August
1943. Sketch by A.N. Formozov, about 2/3 natural size.

119

28,
ъ a?

10-72 ст de os
0&5,

100-110 cm

27-30 ст

3-4 о en у i. sie
© - 2] ст =

yt 30-35.cm

“а eddy
@
Quiet gait while ‘Quiet gait a 96 |
©

searching for while
food traveling Trotting gait Leaping gait

Fig. 34. Tracks of raccoon dog during various gaits (from Yu. Popov, 1956).

reminiscent of badgers, but usually wider. A fan of excavated soil (see
above) is typical of the burrow.

Tracks are rounded in form, nearly 5 x 5 cm for the front, and
4.5 х 5 cm for the hind feet. As a rule, the prints of their claws are
generally easily seen. Unlike foxes, the series of tracks of the running
raccoon dog is not straight, but zig-zag. A raccoon dog when fattening
walks slowly, and looks in each pool, hole and under bushes and
stumps; moving in this way, the tracks of the hind paws do not reach
the tracks of the front. Having short legs they leave “drags” [marks]
and “eyes” [holes] on the snow when it is 10 cm deep. In the snow
cover depth of 25—30 cm, the animal leaves a deep furrow in the loose
snow.

Typical “latrines” in the form of heaps of excrement are found in
definite places, usually 10-15 cm from the den or on trails. Unlike
badgers, the raccoon dog does not bury or hide its excrement. Feces are

94 40-60 mm in length and 20-30 diameter and are composed of 2-3
pieces.

120

Usually silent; only in the rutting period, the males yelp and give
a hollow growl. (A.B.).

Practical Significance

The raccoon dog provides thick, long, strong, but coarse fur. When
cage reared, the animals also provided about 100 g of wool, of a quality
somewhat inferior to that of goats. In a series of regions in the Far East,
the meat of raccoon dog is used by the local inhabitants as food.
Analysis of the meat (18% protein, 3% fat; more than 4186.8
joules/kg'’) reveals its high nutritious qualities (Korneev, 1954).

Data about the size of the catch in the past are almost lacking. In
the eighties of the past century, the trade of raccoon dog in the Primor’e
and Ussuri territories was “quite developed” (Silant’ev, 1898) and
occupied an evident place in the fur trade (Nadarov, 1888). The world
harvest in its entire range, including Japan, North and South Korea and
China in 1907—1910 comprised 260—300 thousand pelts (Kaplin and
others, 1955); of which, judging by indirect data, about 5—8%, i.e. 15—
20 thousand pelts were obtained from Russia. These figures are,
apparently, exaggerated, for the production of the whole fur
industry in the Russian Far East was estimated at about 40 thousand
rubles, and hunting in these regions included sable, red fox, otter,
Kolonok, wolverine, bear, wolf, tiger and others. Even if we suppose
that the raccoon dog furs constituted 50% of this value (which is hardly
probable), then, if the price of raccoon dog fur is 2 rubles, their number
would not exceed 10,000. Probably, the total catch was 5—6 thousand
pelts.

In the 30’s, when lands were considerably opened up, about 12
thousand raccoon dogs were caught.

In the regions of acclimatization licensed trade started in 1948—
1950, and the licensed restrictions were removed in 1953—1955. After
the start of the trade, the number of animals harvested sharply in-
creased, and from 1953 to 1961, it fluctuated within 30—70 thousand
pelts. For example, in 1955, about 41 thousand were obtained, and in
1961—66 thousand. In the latter year, of these about 10 thousand were
obtained from natural habitat in the Far East, and 56 thousand from
places of acclimatization. Of these 56 thousand, Byelorussia provided
6.5 thousand pelts, the Ukraine—about 5 thousand, Latvia, Lithuania

' According to international energy system (SI).

121

and Krasnodarsk territory—4 thousand each, Kalininsk district—3.7
thousand, Pskovsk—2.7 thousand, Astrakhansk—2.3 thousand,
Vologodsk, Moskovsk, Leningradsk, Novgorodsk, Smolensk, and
Yaroslavsk districts, Azerbaidzhan, Estonia, and Dagestan—from | to
2 thousand pelts each. In the remaining districts, territories and repub-
lics, the annual catch was less than one thousand pelts.

Everywhere the raccoon dog is hunted from November until the
snow is deep, i.e. usually until February. In the Far East, the main way
of hunting the raccoon dog is by tracking with laikas or mongrel dogs
at night. In the past century, the Gol’dy* and Orocheny* also hunted
by night with dogs on whose necks were fastened small bells. Less
frequently, they hunted with different “devices” (Nadarov, 1888 and
others). In places of acclimatization, methods of hunting raccoon dogs
are also simple. In the majority of cases, it is hunted incidentally while
hunting other fur animals or specifically harvested with the help of
dogs. With dogs, 80-90% of the animals, on average, are caught; with
guns, 8—10%; with traps, 5—7%. The dog quickly follows the trail and
overtakes the animal, and if it does not flee into a burrow, the dog
strangles it or worries it until the hunter arrives. Traps for the raccoon
dogs are put at burrows, along the shores of water bodies, and around
marshes and ponds.

Cage rearing of raccoon dogs started in the Far East in 1928
(Firsov, 1929). In 1934 there were 15 State farms where the animal
was raised. Collective fur farms in the first year of their existence
selected the raccoon dog as their principal object.

Later, the captive population in the collective fur farms began to
decrease because the raccoon dog requires nearly the same food as
needed by the silver fox. By the end of the 30’s and the beginning of
the 40’s , the breeding of raccoon dog was broadly developed only in
the collective farms of the Ukraine and from 1945 on, this type of fur
husbandry started to decrease (П’та, 1952 and others).

In the 30’s work began on acclimatizing this animal in the Euro-
pean part of USSR, in the Caucasus, and in Siberia. From 1928 to 1958
about 10 thousand individuals were introduced in 76 districts, territo-
ries and republics. The first attempts at transplantation started in the
Primor’e territory where the raccoon dog was brought to some islands
in the Japanese sea. In 1934, it was introduced into the Altai, northern
Caucasus, in Armenia, Kirgizia, Tatariya, Kalininsk, Penzensk and

*Indigenous tribes in the Far East—Sci. Ed.

122

Orenburgsk districts. In the following year—into Leningradsk,
Murmansk, and Novosibirsk districts, Bashkiriya and other regions.
The raccoon dog was most intensively transplanted in 1936 when more
than 1,200 beasts were introduced, and then from 1948 to 1953, when
500—800 individuals were introduced every year in tens of districts
(Shaposhnikov, 1956; Lavrov, 1957 and others).

The acclimatization of raccoon dogs in Irkutsk and Novosibirsk
districts, in Trans-Baikaliya and the Altai did not give results due to
severe winters and inadequate diet. The raccoon dogs acclimatized
poorly in the montane regions of the Caucasus, Trans-Caucasus, Middle
Asia and in Moldavia (Meladze, 1947; Yanushevich, 1956; Arens,
1957; Trushchalova, 1959 and others). Success followed the introduc-
tion of raccoon dog into the European part of USSR, especially in the
northwest (Pribaltic, Byelorussia, Kalininsk, Novgorodsk, Pskovsk,
Smolensk districts), in the central regions (Moskovsk, Yaroslavsk,
Vologodsk, Gorkovsk, Vladimirsk, Ryazansk and other districts), as
well as in the chernozem [black soil] belt (Voronezhsk, Tambovsk,
Kursk districts), the lower Volga region and the level parts of the
northern Caucasus and Dagestan. In the Ukraine, the greatest number
of raccoon dog was established in Poltavsk, Khersonsk and Lugansk
districts.

The trade quality of fur of the raccoon dog acclimatized in the
northwest was somewhat improved. Thus, in Kalininsk district (Morozov,
1955; Sorokin, 1956) raccoon dog fur became somewhat denser and
softer. The length of the guard and top hairs increased by 7.96% on the
average, that of the under fur, by 5.3%. The thickness of the guard and
top hair decreased by 3.41%. The density of the fur increased by
11.3%. The fur color became somewhat darker; the darkest “black-
brown” color in Kalininsk district is found in 8% of the beasts against
3% in their homeland* (Morozov, 1951, 1955; Tserevitinov, 1953).

The importance of the raccoon dog in the hunting economy of a
series of regions is not yet sufficiently clear. Thus, the importance of
this animal is not clear in the Far East. The raccoon dog causes appre-
ciable damage to the pheasant by destroying its nests; this is hardly
compensated by the benefit gained from its destruction of harmful
rodents and insects. In regions of acclimatization the harmful activity
of the raccoon dog in the hunting economy, agriculture and also public
health is obvious. Cases of nest destruction and eating of birds and

*Probably refers to indigenous animals in the Far East—Sci. Ed.

123

nestlings of waterfowl and marsh birds are numerous (Korneev, 1954).
The harm caused by the raccoon dog in floodlands and shores of es-
tuaries is especially sensitive, since, in spring, it feeds entirely on the
eggs and nestlings of waterfowl. In the floodlands of the Oka river,
raccoon dogs destroy a great number of wild fowl; about 46% of its
food consists of birds (Borodin, 1951). In the Voronezh preserve, birds
constitute 48.6% of the food of the raccoon dog litters inhabiting the
floodlands. Moreover, the raccoon dog destroys a considerable number
of birds (45%) during the time of spring migration (Ivanov, 1962). In
Lithuania, birds constitute 15-20% of raccoon dog food and this car-
nivore causes noticeable damage to waterfowl and forest game (Prusaite,
1960, 1961). The raccoon dog frequently destroys muskrat houses and
eats their young (Korneev, 1954; Kritskaya, 1961).

In the Ukraine and in many other southern regions, the raccoon dog
causes harm to kitchen garden and melon cultivation, vineyards and
corn seedlings (Korneev, 1954, and others). Evidently the raccoon dog
plays a negative role as a carrier of rabies, piroplasmosis and scabies
(see above).

A series of authors (Obtemperanskii, 1956; Yu. Popov, 1956;
Sorokin, 1957; Samusenko and Golodushko, 1961), on the basis of the
fact that rodents and insects constitute the main food of raccoon dog
while birds are of secondary importance in its food, consider the harm
caused by these beasts to be negligible. However, the destruction of up
to 90% of the broods of waterfowl (Korneev, 1954) or of a consider-
able number of the broods of forest game and small passerine birds is
hardly compensated by the destruction of mouse-like rodents in the
warm period of year. Undoubtedly, in a series of regions, the harm
caused by the raccoon dog may be relatively small, and in others very
great, and this requires concrete analysis; however, as a whole, damage
by the raccoon dog to the hunting economy is evident.

In spite of the fact that as a result of acclimatization, the prepara-
tion of raccoon dog pelts grew 4—6 times and in a series of districts and
republics its specific contribution to fur production constitutes up to
10-30%, the interests of the protection of nature and sport hunting
economy require the destruction of raccoon dog in places where it has
become acclimatized (A.B.).

Эй

124
GENUS ОЕ WOLVES
Genus Canis Linnaeus, 1758

1758. Canis. Linnaeus. Syst. Naturae. Ed. X, I, p. 38. Canis familiaris
= Canis lupus L. (domestic dog)!.

1816. Thos. Oken. Lehrb. d. Naturg., 3, Th. 2, p. 1037. Thos vulgaris
Oken = Canis aureus Linnaeus.

1816. Lupus. Oken. Ibid., p. 1039. Canis lupus Linnaeus.

1857. Vulpicanis. Blainville. Ann. Sc. Nat. Paris, Zool., 8; pt. 2, p.
279. Canis aureus Linnaeus.

1839. Sacalius. H. Smith. Jardine’s Naturalists Library, Mamm., 25,
p. 214. Sacalius aureus = Canis aureus Linnaeus.

1841. Oxygous. Hodgson. Calcutta Journ. Nat. Hist., 2, p. 213. Canis
aureus Linnaeus.

Large species, but in part of moderate size.

Skull broad and massive, with moderately diverging zygomatic
arches, high (height in the occipital region more than 1/3 of condylobasal
length), in adults and old animals well-developed crests, in particular,
a large sagittal crest. Facial parts of the skull relatively weakly elon-
gated, massive, quite high, and always longer than the cranial parts
(distance from the posterior margin of the infraorbital foramen to the
posterior margin of the canine alveolus is less than or equal to the skull
width above the canines; distance from the line joining the tips of the
supraorbital processes to the anterior end of the premaxillae is greater
than that to the upper part of the occipital condyle). Interorbital (fron-
tal) region convex with large cavities (sinuses) and is clearly and quite
abruptly elevated above the line of the dorsal profile of rostrum. Su-
praorbital processes, which also have cavities, are not large in area, but
massive, and their upper surface is convex, the tip summit bent some-
what downwards, and without sharp borders (edges). Posterior margin
of tooth row extends to the level of the anterior margin of the
interpterygoid fossa. Deep notch on posterior lower margin of lower
jaw anterior to angular process absent.

4: 2
Dental formula в. Teeth large and massive,

'As was shown (see vol. I, p. 540), in this book domestic forms are excluded from
the nomenclature. The wolf, С. /upus, must therefore be considered the generic type, not
only as the most typical species of the genus but also as the ancestral form of “C.
familiaris.”

125

carnassial teeth strongly manifested, palatal length about 2.5 times
length of upper carnassial tooth (P*) and cheek teeth. Canines slightly
curved, massive, broad at base and relatively short. When jaws are
closed, tips of upper canines do not reach middle of mandible, while
tips of lower [canines] do not, or only just, reach margins of alveoli of
upper canines. Tips of upper canines directed straight downwards; no
~ pronounced longitudinal sharp ridge on posterior side of canine, surface
of canine smooth. Cutting edges of incisors with small accessory lateral
cusps (trilobed).

In general appearance, the animals are long-legged, with notice-
ably elongated trunk; tail is fluffy, of moderate length—aup to two-thirds
length of trunk with the head (often less) and not more than twice as
long as hind limbs. When the animal is standing, its end is usually not
lower or only slightly lower than the tarsal joint, but it never reaches
the ground.

Head notably elongated with somewhat pointed muzzle or else
blunter or fairly broad with broad forehead. Ears relatively short with
pointed tips. On sole of hind foot, behind four naked digital pads is
situated a larger fifth. Pads of digits III and IV of fore limbs either
separate or united (jackal, Fig. 17).

Fur quite coarse. Color unimorphic (dimorphic only in some races
of wolf, С. шриз, from southern North Атепса—“С. niger’), grayish,
yellowish, and reddish tones mixed with black hairs, sometimes with
noticeable black dorsal saddle; adult animals never exclusively mono-
tone (except black phase of “С. niger”). Violaceous* gland present,
but weakly developed. Teats usually 5 pairs.

Sharp sexual dimorphism absent, but females are usually some-
what smaller than males. Age dimorphism occurs, and seasonal
[dimorphism] is well marked in some species, but manifested almost
only in density and length of fur. Two molts per year.

Species of the genus are very close to each other and homogeneous,
representing the wolf type—the true gray wolf (С. /upus) and its small
copy—the jackal. The large forms of the wolf have the greatest dimensions
(see family characteristics, and below—description of wolf), the smallest
jackals are not more than 80-90 cm in body length, usually about 70—75
cm, and in weight not more than 10—12 kg, usually about 6~7 kg.

Species of the genus are found in all landscapes from tundra to
lowlands of hot deserts, and high mountains. They prefer regions with

*Also called supracaudal—Sci. Ed.

‘Gouday “D'A) (.08uip 1иво,,) O8uIp эц} Jo э8иел oY} SoUITINO е|елззпу SuUIpUNOUNs sul] OY] "1 51420) зэлом JO пи ay} JO э8ие1 рэзопцзиооэ “SE ‘SI 86

ast

)

2. é
= ie ae к
О

$ У ео =
у a
4 SY
ae у
<. ~Y

127

more or less open areas and abundant wild or domestic hoofed mam-
mals. They are sedentary, [but] individual animals sometimes roam
widely and settle in distant areas. Monogamous animals; moreover,
some form close pairs for several years. 10—13 pups, usually less.
Young usually delivered in open den. Predators, feeding mainly on the
meat of mammals, frequently large ones (ungulates), in part on carrion
and other vertebrates, exceptionally on invertebrates, and at times even
plants. Except during breeding, they usually live and hunt in small
groups. In some species, internal relationships in the group (rank, etc.)
are sharply marked (wolves), in others, quite weakly (jackals).

This genus is very widely distributed, extending in the north to the
northernmost points of land (83°20' М. Lat—not found on floating ice);
also found in equatorial regions. In the New World, it occupies the
entire continent of North America (including the entire Arctic archi-
pelago) and in the south, it extends to Costa Rica inclusive. It is absent
on the Queen Charlotte Islands and islands of the Bering Sea. In
Greenland, it occupies the northern, eastern and western coasts, south-
ward nearly to 70° М. Lat.

In the Old World, its range occupies all of Africa (excluding Mada-
gascar) and all of Eurasia southward to southern China, Siam [Thailand],
Burma, Assam and Ceylon inclusive. In the north, the range includes the
Kolguev Islands [Barents Sea], south island of Novaya Zemlya, Belyi
[island] adjacent to the Taimyr coast, as well as Lyakhovskii Islands and
the New Siberian archipelago. They are also found in Kamchatka, Sakhalin
and the Japanese islands. They are absent on Iceland, Spitsbergen, Franz
Josef Land, Severnaya Zemlya, Wrangel Island, the Commander Islands,
the majority of the Kuril islands and Shantar Island, Taiwan, Hainan, and
the Malacca and Malaya archipelago.’ At the present time, it is extermi-
nated in a series of territories of North America, in the greater part of West
and Central Europe, in part of Japan and in several other places. This genus
represents a highly adaptable and viable group, and its range has decreased
only in the regions of dense human population and where it has been
directly persecuted to extinction.

The genus appeared in the upper Pliocene of North America; in
Eurasia, in the Pleistocene (see characteristics of genus Vulpes).

Particular generic or subgeneric names have been proposed for
almost every species included in the genus of contents accepted here.
This occurred during the period of narrow interpretation of the species

°A feral domestic dog—‘Canis dingo” —is found in a completely wild condition in
Australia (except Tasmania).

100

128

concept, when many of the races were considered to be separate spe-
cies, and these [genera or subgenera] could be group names. At the
present time, there is no basis for dividing the genus into several
subgenera, and it is senseless to separate, within it, such subgenera as
Canis for the wolf and Thos, Schaeffia, Lupulella for the various forms
of jackals. The differences between these “subgenera” are very insig-
nificant; moreover, in the present species concept, only one species is
included in each subgenus. Division of the genus into two subgenera:
Canis and Thos, i.e. wolf and jackal is also not justified. In some
respects, the species of the genus themselves represent the fullest and
most characteristic expression of the family type.

This genus is most closely related to the group Simenia (Abyssinian
wolf) and Vulpes (foxes) and especially to Alopex (arctic fox). This
latter represents, in craniological respects, a quite clear transition and
connecting link between Canis and Vulpes. In the past, all these groups,
especially Canis, Vulpes and Alopex, were usually grouped in one genus
or were considered as subgenera only. Later, however, all authors es-
sentially divided these groups generically or, at least, separated Canis
from Vulpes, united Alopex with the latter group, and separated Simenia.
Most recently, it was again proposed (Haltenorth, 1958) that all the
mentioned groups including a series of others be considered members of
one enlarged genus—Canis (see characteristics of the family). In this
account, the described groups occupy subgeneric positions. This point of
view deserves consideration, but requires broader argument.

In the genus are 5 species: 2 African—C. adustus (striped jackal)
and C. mesomelas (shabrack [= black-backed] jackal), 1 North Ameri-
can—C. latrans (coyote), 1 Eurasiatic—African— С. aureus (jackal)
and 1 Eurasiatic-North American—C. lupus (wolf).? The species of
this genus constitute about 17% of the species of the family.

One species—the wolf—is found both in the wild and in the domestic
state as the domestic dog (“Canis familiaris”). The dog—the most strongly
altered form as compared to its wild ancestor—is generally the most strongly
divergent and diverse of domestic animals (the number of breeds exceeds
200). Nevertheless, in respect to the wolf, there is no reproductive isolation.
The view which was widespread not long ago, concerning the diphyletic
origin of the domestic dog—from wolf and jackal—is now totally dis-
regarded. As in cther domestic animals, the dog has one original species
(“ancestor”). The possibility of the diphyletic origin of the dog is excluded

>The wolf of Florida and the southeastern states adjacent to the Mississippi
[River], is considered by American authors to be a separate species—C. niger.

129

in particular by the difference in the diploid number of chromosomes—78
in wolf and dog and 74 in jackal (Matthey, 1954).

In the USSR, two species are found: wolf, Canis lupus Linnaeus, 1758
and jackal, Canis aureus Linnaeus 1758—about 40% of the species of the
genus and about 0.6% of the number of mammals of the Union.

The range of the genus covers the entire territory of the USSR,
except the above-mentioned northern territories and islands.

The species of the genus are of great importance in the USSR as
pests of livestock and the hunting economy and, to a lesser extent, as
carriers of rabies. Their importance as fur-bearing species is not great
and less than the harm they cause (V.H.).

JACKAL, CHEKALKA
Canis aureus Linnaeus, 1758

1758. Canis aureus Linnaeus. Syst. nat. Ed. X, 1, p. 40. Laristan,
southern Iran.

1835. Canis aureus var. moreotica. Geoffroy. Exped. Scient. de Moree,
Zool., pl. 1. Peloponnesus (Morea), Greece.

1858. Canis aureus typicus oder var. caucasica. Kolenati.
Reiseerinnerungen. Th. I, p. 96, Armenia. (V.H.).

Diagnosis

Adult body length not more than 90 cm, tail length about one-third of
body length. Length of skull less than 200 mm. Arciform line formed
by anterior border of both nasal bones has, in the middle (along suture
between nasal bones), a well-marked projection, directed anteriorly. At
antero-internal sides of auditory bullae, swellings in form of ridges on
basioccipital absent. Cingulum on outer surface of first upper molar
well marked, broad and continuous (Figs. 15, 16). (V.H.).

Description

In its general appearance, the jackal is very similar to the wolf, but
much smaller in size, lighter in weight, with shorter legs and with a
somewhat more elongated trunk and shorter tail. Its end just reaches the
heel or a bit below it. As in the wolf, the tail always droops. The head

101

130

is lighter, not so “foreheaded”*, and the muzzle is narrower and more
pointed. On the whole, the jackal much resembles a small mongrel dog.
Fur is coarse and stiff. Iris is light or dark brownish. Teats 5 pairs.

General color of winter fur is dirty reddish-gray, strongly high-
lighted with blackish due to the black tips of many guard hairs, or a
brighter, rusty-reddish color. Anterior рай of muzzle, circumorbital
region and forehead are ocherous-rusty-reddish. Above each eye, a
blackish stripe is present. Margins of lips and lower cheeks are dirty
white. Upper part of forehead and occiput are ocherous. Back of ears
is pale-rusty; ears are covered internally with dirty whitish hairs. Chin
and throat are whitish, with a dirty tint.

Black guard hairs are especially developed on the back, but less so
on the sides, and general color here is brighter and clearer. Belly is
whitish along the midline, and in the axillary region and groin the color
is mixed with a reddish tint. Extremities are ocherous-red, with internal
surfaces of lighter color. Tail is gray with an ocherous tint and a
strongly defined dark color on dorsal side and at tip. General color
intensity and the degree of development of darkening is quite strongly
variable individually, and both comparatively dull jackals as well as
those with very bright colors are encountered.

CN \ —
ih ra:
Wee:

о жа ’
4
= is у: м
7 GN Ee
Е ЗА 9
я 5
"0

Fig. 36. Jackal, Canis aureus L. Sketch by A.N. Komarov.

*Russian word is “Jobastaya”, referring to the more expanded frontal region of the
wolf compared to the jackal—Sci. Ed.

103

131

Summer fur is more sparse, coarse and short and has same color
as that of winter, but brighter with dark tint less developed. Sexual
differences in color are absent. Pelage of newborns is very soft and its
color varies from light-gray to dark-brown. This pelage remains on the
animal for about one month and then the first adult coat starts to grow
and is developed by August. It is very similar to that of adults. Geo-
graphic variation in color is fairly noticeable.

The skull is similar to that of the wolf in its main features and in
general appearance, but is smaller and less massive. The nasal region
is lower, and the facial shorter (its length is nearly equal to that of the
cranial region), dorsal profile of the facial region is nearly straight and
depression in middle of nasal bones is very weak. In the region of the
carnassial teeth, the skull is not broadened and its outline, beginning
with the line of the zygomatic arches, converges anteriorly in a quite
regular wedge (on looking up at the skull from below, a sharp projec-
tion in front of the carnassial tooth is not seen). Sagittal and occipital
crests are well developed, but weaker than in the wolf. Supraorbital
processes are large, but relatively less massive than in the wolf. Be-
tween the frontals and between the nasals, along the corresponding
sutures, a shallow longitudinal depression is present. On the line of the
anterior margins of the nasal bones (at suture between 2 nasal bones)
there is a distinct projection (Fig. 15). At the anterior internal borders
of the auditory bullae at the basioccipital, the bulla is not bordered by
bony ridges.

Canines are large and strong, but relatively thinner than in the
wolf, and carnassial teeth are relatively weaker. The cingulum on the
outer margin of the first upper cheek teeth is continuous, broad and
well defined throughout its whole extent (Fig. 16).

Sexual differences in the skull are only manifested in the generally
somewhat smaller dimensions of the female. Age variations correspond,
in all principal aspects, with those described in the wolf.

Diploid number of chromosomes—74.

Male body length 71-85 cm (as a rare exception in the species,
may be somewhat more), females average somewhat less.‘

“In regions on the right tributaries of Pyandzh it was shown (Chernyshev, 1958)
that body length of males (14) is 68-104 cm, and of females (17) is 71-100 cm. These
maximal figures are, apparently, exaggerated. Such figures are not given by any of our
authors for either Middle Asiatic or Caucasian animals. Mean body length, drawn from
the tables given by the mentioned authors, is 81.5 cm for males and 80.5 cm for females.
The maximal specimen, of 75 caught in India, Baluchistan and [ran had a body length
of 81 cm (male) and 73.5 cm (female). Six females from Nepal had an average body

132

Measurements of animals from southern Tadzhikistan (region of
the right tributaries of Pyandzh and upper Amu-Dar’ ya) are as follows.
Adult (sexually mature) males: tail length (14) 22—27 cm, length of
hind foot (13) 13.7—19 cm, height of ear (8) 6.7—10 cm. Adult females:
tail length (17) 20-27.5 cm, length of hind foot (17) 9.5—12 cm, height
of ear (15) 7.1-9.1 cm. Height at shoulder (12) of males and females
44.5-50 cm, height at sacrum 40—48 cm.

Weight of males (15) 6300—13,670 g, of females (16) 7000—
11,240 g.

Weight of heart 100—150 g, of liver 185—480 g, total. length of
intestine 335—454 cm, length of caecum 7.5—12 cm (Chernyshev, 1954).

Measurements of male skulls (14 from Caucasus and from Turkestan;
Ognev, 1931; ZMMU*): condylobasal length 147—164 mm, zygomatic
width 79-97 mm, breadth of muzzle above canines 27-32 mm,
interorbital 22.6-30.2 mm. Skull of females (17) from the right bank
of upper Amu-Dar’ ya (Chernyshev, 1958): condylobasal length 123.7—
159.7 mm, zygomatic width 69.9-92.5 mm, length of upper toothrow
64.5—87.5 mm; interorbital width 19.4-30.1.

Measurements of jackals within the USSR do not show clear geo-
graphic variation. (V.H.)

Systematic Position

The jackal must be considered as a somewhat less specialized form of
the genus than the wolf. This is manifested in the relatively short facial
portion in the somewhat weaker tooth row, with which, in particular,
the above-mentioned “wedge-shaped” skull and its relatively small width
in the region of carnassial teeth is connected, and in the less developed
crests. This is, apparently, connected with its diet of small prey—irds,
rodents, lower vertebrates, insects, other animals and carrion. The jackal

length of 73 cm, 12 females from Palanpur and Sind—69 cm (Pocock, 1941). It is
generally considered that Indian animals have a body length of 61—76 cm (Prater, 1947).
Maximal Balkan jackals had a body length of 85.5 (male) and 81 cm (female; Atanasov,
1953). Our jackals are, apparently, slightly larger than the Indian ones, but not by much.
It is considered, under the circumstances, a mistake, that the skull dimensions given for
Tadzhikistan, do not differ from generally known ones. One foreign author (Van den
Brink, 1958), who did not refer to the source, gave still more exaggerated dimensions
for the European jackal: 85—105 cm, and another (Marches [et al.], 1954) gives a body
length of 104 cm for a male and 100 cm for a female from Romania. A weight for jackal
of “more than 16 kg” (P. Manteifel; Chernyshev, 1958) is also exaggerated.
*Zoological Museum, Moscow State University.

133

102 Fig. 37. Skull of jackal, Canis aureus Г.

104

134

is a “scavenger” in no lesser degree than it is a predator. This is,
apparently, the significance of the somewhat smaller general dimen-

. sions of the animal (V.H.).

Geographic Distribution

Southern Asia and Near East, southeastern Europe, northern half of
Africa.

Geographic Range in the Soviet Union

This constitutes a considerable part of the species range—its northern
border is connected with the southern part of the country—Caucasus,
Middle Asia and Moldavia. It consists of several parts within the Soviet
Union separated from each other which unite beyond our borders.

In the Caucasus, the range of the jackal is connected with the
plains and foothills, in places with low and even middle montane belts,
and having quite complicated contour. In the western half of the coun-
try, the jackal is distributed in a narrow strip along the entire Black Sea
coast from Novorossiisk to Batumi. Here, the animals are not usually
distributed in mountains higher than 400—500 m above sea level. The
range widens somewhat along the Rion Basin and extends somewhat to
the east of Kutais’.

In the beginning and middle of the past century, the jackal was,
apparently, distributed (perhaps in part as vagrants, however this was
not very rare) along the eastern shore of the Azov Sea to the lower Don
and even to Taganrog. Old reports about the occurrence of the de-
scribed species in Crimea are not worthy of attention.

In the eastern half of the Caucasus, beyond the Main Range, the
jackal is distributed on the plains and foothills and, as a rule, is not
found on mountains higher than 800—900 m above sea level. Only in
Zakatal, in autumn, it may climb up to 1000 m searching for fruits, and
everywhere in Talysh, to a height of 1700 m (the highest occurrence
known in the USSR). In the Trans-Caucasus, the jackal is distributed
from Talysh and the Lenkoran lowland southward to the Apsheron
peninsula and the foothills of the Great Caucasus on the north. Here,
it extends to Zakatal and a little more to the north. In the west, the
jackal is distributed along the lowland places of the Iora, Alazan’ and
Kura basins, to the Tbilisi meridian and a little to the west (Mtskhet).
Southward, the foothills of the Little Caucasus constitute the western
border of the eastern Trans-Caucasian part of the range.

106

135

Along the valleys of various rivers, the jackal penetrates in some
places in a minor way into the mountains. Along the Araks valley, the
range extends in a narrow strip westward to Oktemberyansk region
(44° Е. Long., a little east of Erevan). In the Araks, the animal does
not ascend higher than 500-850 т above sea level.

From the Apsheron peninsula, the range of jackal extends, in a
narrow strip, along the Caspian Sea northward to the mouth of the
Terek and Kuma and to the Beryuzyak peninsula. In part of the range
northward to Makhachkala, the jackal is distributed along the sea shore,
generally not further than 20-30 km from it. Beginning only at
Makhachkala, it goes deeper into the country along the foothills and
river valleys. To the west of Makhachkala, the jackal is distributed in
a narrow strip along the foothill belt to Khasav”yurt and a little west-
ward. It goes west along the Terek to Shelkova (Shelkozavodsk) and
Parabochevsk forest and reaches Mozdok, and along the Sunzha, to
Groznov. It is not found along the Kuma river upstream from its mouth.

In the past, jackals (apparently, mainly along the Caspian coast),
reached the delta of the Volga (30’s and 40’s of the nineteenth century)
and were also found in “Don steppes” and still in the middle of the past
century along the Ergeni river. They also inhabited Stavropol’ at that
time. It is now impossible to decide whether the places of occurrence
along the Ergeni and Don were connected with the Pri-Caspian region
of occurrence of the species, or with the Pri-Azov region. It is probable
that all these were one region.

Of the Caspian Sea islands, the jackal lives only on Sar Island at |
Lenkoran’. It is absent on other even larger (Chechen’) islands as well
as on Agrakhansk peninsula (spit).

In the last ten years, to a considerable degree in connection with
the drying of the Caspian Sea and the disappearance of reed beds, the
range of the jackal in the northern Caucasus has contracted. In particu-
lar, it does not reach Kuma.

In the northern Caucasus, chiefly at the beginning of the present
century, fairly long intrusions are known—to Gunib in the heart of the
mountains of Dagestan, to Stavropol’, to Psebai (south of Armavir), to
the mouth of the Kuban’ river and even into the mouth of the Don.
Wanderers in the eastern parts apparently came from the Kizlyar ге-
gion of occurrence of the animal, and in the west, from the Black Sea
coast. Jackal intrusion even to Tambov (Ognev, 1931) is known, prob-
ably from the Caucasian region of occurrence of the animal.

The distribution areas of the jackal in the western and eastern
halves of the Caucasus are, in essence, separated from each other, but

4 = an
ke _ < м

сх

es а.
nord
5 <
<Я

ду
K

aS РАЙ т
У:

ALS
SEC) Я

s. V.G. Heptner.

1 distant intrusion

L. in the USSR. Points beyond the limits of the range refer to severa

105 Fig. 38. Range of the jackal, Canis aureus

137

in the middle of the Trans-Caucasus they are very near to each other,
separated only by the Suramsk range and the adjacent regions—a total
of 100 or 100+ km. Animals occur only as intruders and their move-
ments, apparently, reflect some kind of connection that is or was realized
between the two described divisions of the range. The present union of
the above-mentioned parts of the distribution area applies, however,
only farther the south, outside the boundaries of the USSR.”

The Middle Asiatic region of jackal occurrence within the Soviet
Union is completely isolated from that of the Caucasus. It has quite a
complicated form. This is explained by the fact that the jackal, in its
distribution here, is associated with river valleys and cultivated lands,
resolutely avoids deserts, and does not go high into the mountains. It
is, in part, divided into separated sections, isolated from each other. In
the west, in Turkmentya, the jackal is found along the Atrek, from
Chikishlyar westward to the mountains. Along the river valleys of the
Atrek, Sumbar and Chandyr, it penetrates into the western parts of the
Kopet-Dag (at least to Kara-Kala on the Sumbar and Yarty-Kala on
the Chandyr), but it does not get into the heart of the Kopet-Dag
mountains. Its range envelops the Kopet-Dag from the west (?) and
north as a narrow strip along the foothill plains and foothills, and
extends to Tedzhen’®. The jackal occupies the entire Tedzhen valley to
its extreme lower reaches and all the valley of the Murgab with its
upper tributaries (Kushka, perhaps Egri-Gek). In the desert between the
Tedzhen and Murgab, and between the Murgab and Amu-Dar’ya, the
jackal is absent. The union of these sections is realized in Iran and
Afghanistan.

Along the Amu-Dar’ya, the jackal is distributed from the Chubek
(on the river, somewhat east of the mouth of the Kizyl-su, nearly
directly south of Kulyab) to the delta which occupies the whole region.
In the basin of the upper Amu-Dar’ya, the jackal is distributed along
the Kizyl-su and Yakh-su northward, nearly to the latitude of Kulyab
or somewhat farther north; along the Vakhsh, it goes up to Kurgan-
Tyub; along the Kafirnigan, nearly to Dushanbe or a little to
the northeast, to the mouth of the Varzob; along the Surkhan-Dar’ya

‘The distribution in the Caucasus according to materials of Nordmann, 1840;
Kornilov, 1859; V.V. Shch., 1887; Alferaki, 1910; Satunin, 1915; Ognev, 1931; Heptner
and Formozov, 1941; Dal’, 1954; Heptner, Turova and Tsalkin, 1950; Kirikov, 1959 and
the unpublished material of V.G. Heptner.

Тре distribution of the jackal in this region is poorly delimited. Apparently, the
mentioned strip is not continuous. In sand and loess deserts, the jackal is absent. It is
also absent in the Great Balkhan and hardly reaches the West Uzboi.

107

138

upstream, at least to Regar. In this way, the range includes the
entire Gissar valley and the foothills of Gissar range eastward to
Ordzhonikidzeabad and almost to Ramit. Between the Surkhan-Dar’ ya
[river] and Kugitangtau and Baisun[tau] mountains, the range protrudes
northward from the Amu-Dar’ya, extending to the city of Chirabad. In
all the described sections on the right bank of the Amu-Dar’ya, the
jackal does not ascend higher than 1000—1100 m above sea level and
the boundary of its distribution envelops, as it were, the higher regions
such as Babadag and other massifs lying along watersheds, and forms
here a fairly complicated pattern.

Fig. 39. Reconstructed species range of the jackal, Canis aureus L. V.G. Heptner.

107

108

139

The range of the jackal further occupies the entire area of culti-
vated land along the Kashka-Dar’ ya (Karski, Chakhrizyabs). This region,
apparently, is more or less isolated from the previously mentioned
Baisuntau mountains. The jackal exists everywhere in the Zeravshan
basin (Bukhara and Samarkand oases), penetrating upwards along the
river to the mountains, to Pendzhikent, and perhaps somewhat higher.
The jackal inhabits the cultivated strip and adjacent places between
Samarkand and Tashkent and is widely distributed in Tashkent oasis,
eastward to the mountains. Along the Syr-Dar’ya, it penetrates to the
western parts of the Fergana valley and to the Farkhandsk reservoir and
Kanibadam (for the eastern parts of the valley, almost nothing is known).
Downstream along the Syr-Dar’ya, the jackal is normally distributed
to the mouth of the Aris. East of the Syr-Dar’ ya, at the northern limits
of its distribution, 1t extends to Shimkent, Lenger, Georgievk’ and
Boroldaisk mountain, and to the sources of Bugun river lying approxi-
mately 150 km north of Chimkent.

In Middle Asia, distant intrusions of the jackal are also known and
moreover they take place partly in years of severe cattle losses (con-
tinuous snow cover)*. In these years, intrusions are more frequent and
extend particularly far. Intrusions of jackals may occur in the lower
Syr-Dar’ya, reaching Kazalinsk; on the Sary-su river; on the Chu river
to its lower reaches; to the northwestern edge of the Betpak-Dala desert
and almost reaching even to the city of Karaganda. To the north intru-
sions are known to the Ustyurt (Kos-Bulak mountain), on the
northwestern shore of the Aral Sea (Kintekchi), in the Asmantai-Matai
sands at the northern base of the Ustyurt, at Emba. Jackals wandered
Turgai and Sary-Kopa and even to Orenburg (see below for details)’.

The third region in which the jackal is known in the USSR is
Moldavia. Here, the animals are rare, appearing as vagrants, and not
coming this far every year, in the southern and southwestern regions.
They enter from Romania (Kuznetsov, 1952). The appearance of the
vagrant jackals in the Trans-Carpathian region cannot be excluded.

*The Mongolian word dzhut refers to persistent continuous snow cover that pré-
vents ungulates from feeding—Sci. Ed.

’Range in Middle Asia according to Ognev, 1931; Laptev, 1934; Flerov and Gromov,
1934; Flerov, 1935; Shestoperov, 1936, 1936a, 1936b; Gromov, 1937; Leviev, 1939;
Salikhbaev, 1939; Sludskii, 1939; Sultanov, 1939; Kuznetsov, 1948; Bazhanov, 1951;
Shukurov, 1951; Afanas’ev et al., 1953; Chernyshev, 1954, 1958; Heptner, 1956 and
others and according to the unpublished material of V.G. Heptner.

140
Geographic Range outside the Soviet Union

The range in Asia includes southwest Siam, Burma, Assam, Bengal, all
of India; in the north, it includes Bhutan, Sikkim, Nepal (in Himalaya
to a height over 3000 m above sea level), Ceylon, Baluchistan and
Afghanistan (except the montane areas of the Hindu Kush), Iran, Asia
Minor and the entire Arabian peninsula. In Africa, the range includes
Kenya, Ethiopia, Somalia, Sudan, Egypt, Libya, Tunisia, Algeria,
Morocco, Rio de Oro [Spanish West Africa, now Western Sahara], and
Senegal. In Europe, the jackal is found in Greece, Albania, Yugoslavia
and Bulgaria. As a vagrant known in Hungary (in the north to Komitata
Khevesh near Tissa northeast of Budapest) and in Romania (very rare,
only in the southwestern part of the country at the Danube south of
Kraiov)®. (V.H.).

Geographic Variation .

Throughout a broad area covering the country, with extremely different
natural conditions, geographic variation of the jackal is quite well
defined, though less than what is accepted. Features of size as well as
color vary. In the USSR, there occur two well-differentiated forms.

1—Turkestan jackal, С. a. aureus Linnaeus, 1758.

General color is relatively pale with sandy tones predominating.
The fur is relatively soft. Dimensions large.

Middle Asia. Outside the USSR—Afghanistan, Iran, Iraq, Arabian
peninsula, Baluchistan, northwestern India (Kutch, Sind, Gujarat). That
animals inhabiting southwestern Turkmeniya are not completely iden-
tical to those living east of the Amu-Dar’ya is not excluded.

2—Caucasian jackal, С. a. moreotica Geoffroy, 1835 (syn. typicus,
caucasicus).

General color is bright and saturated, with strongly pronounced dark
blackish tones on the back. Thighs and upper parts of legs, ears and
forehead are bright, reddish chestnut. Fur is coarse. Size somewhat less.

Caucasus, Moldavia. Outside the USSR—Southeastern Europe, Asia
Minor.

SCalinescu, 1930; Atanasov, 1953. Judging by the catch in Moldavia, one may
consider that intrusions occur also in other parts of Romania and take place more
frequently than is known.

109

141

In territories lying beyond our borders, the following forms are usually
accepted: С. а. syriacus Hempr. et Ehrenb., 1833 (Syria, Palestine); С.
a. lupaster Hempr. et Ehrenb., 1833 (Libya, Egypt, Palestine); C. a.
indicus Hodgs., 1833 (Nepal, Sikkim, Bhutan, Assam, Burma, Siam);
С. a. пана Wrought., 1916 (south part of India); С. a. lanka Wrought.,
1916 (Ceylon); C. a. algiriensis Wagn., 1841 (Tripoli, Algeria, Tunisia);
C. a. maroccanus Cabrera, 1921 (Morocco); C. a. soudanicus Thos.,
1903 (Egypt, Nubian desert, Kordofan, Sudan). Some of these forms
are highly doubtful. Sometimes the accepted (Ellerman and Morrison-
Scott, 1951) C. a. cruesemanni Matschie, 1900, described according
to a living specimen by an author of very little reliability, is not taken
into account here. The existence of the name ecsedensis Kretzoi, 1947
(hungaricus Ehlik, minor Mojsis.) given to animals from Hungary is
also not credited. These are, undoubtedly, vagrant specimen of the
Balkan moreoticus; jackals do not live permanently in Hungary.

The statements that there are 20 species of jackals in the Old
World, not including the black-backed (Ognev, 1931), 15 completely
unsubstantiated. The true jackals, together with the black-backed, are
only 3 species (aureus, adustus, mesomelas). The previous conception
was based on binary designation of races, the unnecessary division of
species and subspecies and upon unreviewed synonymy. (V.H.)

Biology

Population. Along the Caucasian coast of the Black Sea, at Novorossiisk,
the jackal is rare; to the south, it is common, while in Georgia it is
abundant. At the mouth of Terek, it is rare; southward, its number
increases, reaching a maximum in southern Azerbaidzhan. In Dagestan,
from 1936 to 1941, an average of 580 jackal furs were prepared yearly.
The jackal is abundant in the Kolkhid, Zakatalo-Ismailinsk, Kurinsk,
Lenkoran and other lowlands in Georgia and Azerbaidzhan. In Arme-
nia and Nakhichevan Autonomous SSR, they are encountered in small
numbers. In the Trans-Caucasus, in the 30’s of the present century, up
to 17 thousand jackals were caught yearly. In the Trans-Caucasus,
jackals are most common in Azerbaidzhan. From 1931 to 1940 an
average of 9400 animals was caught here yearly, in Georgia—2300, in
Armenia—some hundreds (Vereshchagin, 1947).

In Turkmeniya, it is abundant in the valleys of rivers and rare in
the foothills and low mountains, from whence, from time to time, it
penetrates in small numbers to the very edge of the desert. In higher

142

areas deep within the Kopet-Dag, it is absent or rare (V.G. Heptner).
On the lower Atrek, it is not numerous and comparatively few are
captured (106 in 1941; Samorodov, 1953). It is rare in West Uzboi. In
the valleys of the Tedzhen, Murgab, and Amu-Dar’ya rivers, and in the
oasis irrigated by them, it is abundant; and is more rarely met with
along the Sumbar and Kushka. In the mountains of southern Turkmeniya,
it is encountered regularly. It is known in Karabil’, the mountain be-
tween the Kushka and Murgab rivers, and on the Gyaz’-Gyadyk (upper
Tedzhen), but in small numbers (V.G. Heptner). From 1948 to 1958,
from 1400 to 5400 jackals were caught annually in Turkmeniya
(Sapozhenkov, 1960).

In Tadzhikistan, jackals are abundant in the valleys of the Pyandzh,
Vakhsh, and Amu-Dar’ya rivers and their tributaries—the Kafirnigan,
Kyzyl-su, Surkhan-Dar’ya and in the adjacent cultivated lands. In the
offshoots and foothills of the mountains they are rare. It is especially
numerous in the Pyandzh tugais* from Faizabad-Kala to the mouth of
the Vakhsh and along the entire Vakhsh valley where, however, they
avoid sandy deserts. In the settlement of Tigrov Balka in the Pyandzh
valley, in 1946-47, on an area of 600 hectares, about 133—161 jackals
were counted, i.e. up to 0.02 individual per hectare. In the mountains of
the Vakhsh—Kafirnigan interfluve, it is encountered only during migra-
tions. To the west of the Kafirnigan, and to the north of the Amu-Dar’ya
tugais, it is very rare in the foothills of Koikutau and Tuyuntau, inhab-
iting the whole of the cultivated strip of the Gissar valley, and it is also
not rare in the foothills of Gissar range (Flerov, 1935; Chernyshev,
1954, 1958). It is common in the valleys of the upper reaches of the
Syr-Dar’ya and Zeravshan rivers (Chernyshev, 1954). From 1951 to
1958 from 530 to 1400 jackals were caught in Tadzhikistan.

In Uzbekistan, the jackal is numerous in the valleys of rivers
Surkhan-Dar’ya, Kashka-Dar’ya, Zeravshan, Syr-Dar’ya, in the lower
Amu-Dar’ya and in Khorezm and Bukhara oases; it is rare in foothills
and also in the Fergana valley in the floodplains of the Naryn and Syr-
Dar’ya rivers. In Fergana, it is captured occasionally (Malenkov, 1958).
Only in the lower Amu-Dar’ya, in the territory of Karakalpak Autono-
mous SSR, are about 1000 jackal furs prepared yearly (Kostin, 1956).
In the Amu-Dar’ya delta about 200 jackals were estimated in 1948 on
six tugai woodlots (Salikhbaev, 1950). From 1950 to 1958, from 3300
to 6200 individuals were prepared annually in Uzbekistan.

*The Russian ига! refers to a distinctive riparian shrubland found along rivers of
Middle Asia—Sci. Ed.

110

143

It is rare in the Syr-Dar’ya valley in its middle course north of
Kzyl-Orda, but in some years, it is very common from its confluence
with the Arys’ and farther south, and also in the spurs and foothills of
the Talask Alatau in the territory of Chimkentsk district.

Habitat. In the Caucasus and Trans-Caucasus, the jackal is a low-
land dweller, and as a rule does not ascend into the mountains higher
than 600 m above sea level. Only in a few places (Borzhomi), it is
found at a height of 900-1050 m (Dinnik, 1914), and in Armenia at
present climbs to heights of 840 m (Dal’, 1954). It inhabits the coasts
of the Black and Caspian seas and the level valleys of rivers every-
where where there is dense forest, thickets of prickly bushes and
extensive reed floodlands rich in game and small rodents. Where dense
shrubs are absent, jackals are absent. Preferred habitats are impassable
thickets of bushes, consisting of various spiny and climbing plants such
as blackberry, Jerusalem thorn (Paliurus australis), Smilax excelsa,
Clematis sp. and others. In these thickets, the yearly falling leaves and
the dry branches form, at some height from the ground, a unique floor-
ing that divides the thicket into two levels: in the lower, the inhabitants
of these thickets move freely—yjackals, jungle cats, badgers, wild boars,
pheasants and others feeling themselves here to be in comparative
safety (Satunin, 1915). Such habitats serve as shelters for jackals in
southern Dagestan, on the lower Samur. They are nearly inaccessible
to humans, but the animals freely move there along “tunnels” made by
wild boars (Heptner and Formozov, 1941).

The jackal hunts along rivers and canals, on the sea shore and
around lakes, beating down, together with other animals, the trail sys-
tem. In dry years, when lakes and swamps dry out strongly, jackals
concentrate around them, hunting waterfowl, nutria and small rodents.
In severe winters, when water bodies are covered with ice, the carni-
vores gather around them, cross over to the island and settle in the
reedbeds away from the shore and through the course of the entire
period of frost, do not leave the water bodies. In spring, summer and
autumn, 70 jackals and 60 jungle cats were caught in a small section
of the Inkitsk swamps (Georgia) (M. Pavlov, 1953). Jackals live very
close to settlements. One time a jackal lived in the center of Lenkoran
village under a forester’s house.

In Middle Asia and Kazakhstan, the jackal is also a lowland dweller.
Here, the preferred habitats are tugai thickets and reed floodlands in the
floodplains of rivers, bushes and reeds along the canals and aryks,* and

*The Russian arvk is a Turkic loan word for irrigation ditch—Sci. Ed.

111

144

Fig. 40. Habitat of jackal. Outer margin of tugai of tamarisk and turanga, in
the Murgab valley at Imam-baba, Turkmeniya, 22 May 1962. Photograph by
A.A. Sludskii.

also thickets on abandoned irrigated lands. They prefer wild olive-
turanga and tall grass-herb tugais. Here, trees are densely interwoven
with vines and surrounded with sturdy thickets of reeds or plume grass
[Erianthus]. Places where jackals live usually abound in waterfowl,
pheasants, tolai hares, small rodents, frogs and fish. Here it finds good
cover and abundant prey.

In the Gissar and Fergana valleys, jackals do not find hiding places
in the arable fields in the valley, and live in adjacent low hillocks
[called] adyr. In the adyrs it shelters in dry channels, loess caves and
niches, occupying abandoned holes of foxes and porcupines, or digging
its own holes. In Turkmeniya, if there are-not good thickets along the
river, as for example on the Kushka, the jackal also makes a hole found
in the neighboring hillocks (Heptner, 1956).

The jackal willingly settles in oases, where it confines itself to
thickets along irrigation ditches and reservoirs, in gardens, graveyards,
ruins, etc. In Bairam-Ali, on the lower Murgab, jackals colonized and
dug holes in planted groves of white acacia in a very populous place
(V.G. Heptner). In Middle Asia, the jackal is, to a considerable degree,

145

Fig. 41. Biotope of jackal—tugai on the Murgab at Sultan-bent. Turkmeniya,
10 June 1962. Here also the spotted [steppe] cat and wild boar are found.
Photograph by A.A. Sludskii.

a synanthropic animal. It is especially abundant in very old oases, such
as Tedzhen, Pendin or Merv on the Murgab, Bukhara on the Zeravshan,
and Khorezma on the Amu-Dar’ya. In connection with irrigation and
utilization of deserts and the establishment of new oases in the 30’s—
50’s of the present century, rapid colonization of jackals was observed
in the new regions lying in the Vakhsh—Pyandzh interfluve, in the
Golodnaya steppe of Uzbekistan and in other places. A further increase
in the number and dispersion of jackal may be expected.

The jackal avoids waterless deserts, being encountered there only
at their very edge. As a rule, it does not live in the Karakum and
Kyzylkum, but is met with in significant quantities along the borders
of the sandy deserts, adjacent to the Murgab and Amu-Dar’ya oases
(Salikhbaev, 1950; Stal’makova, 1955), where it comes out from the
tugais and thickets in order to hunt gerbils. In Repetek, situated only
70 km away from the Amu-Dar’ya, jackals were not observed
(Stal’makov, 1955). However, in the fall of 1957 and 1958, they
appeared here (Sapozhenkov, 1960). They appear in the Asmantai sands

112

146

in the Ustyurt, migrating from the delta of the Amu-Dar’ya and from
the shore of the Aral Sea. In foothills and mountains they are confined
to tugai shrubs along ravines, and in bushes along rivers and gullies.
In the Himalayas, it ascends somewhat higher than 1000 m above sea
level.

The presence of jackals in one or another region and their choice
of habitats are determined by the abundance and availability of prey,
the presence of water and dense shrubs which serve as shelters both for
the jackals themselves and for the animals which they hunt. It is espe-
cially abundant there where water bodies incur no prolonged freezing
period and where it is possible for waterfowl to overwinter.

Jackal can withstand low temperatures down to —25° [C] and even
—35°, but it is not adapted to live in very snowy lands. In the Trans-
Caucasus, in severe winter with much snow (1949—1950), jackals could
travel only on roads and paths made by humans and large animals
(Vereshchagin, 1951).

Fig. 42. Habitat of jackal on the Murgab. Tugai of turanga and tamarisk at
Sultan-bent, Turkmeniya. 10 June 1962. Photograph by A.A. Sludskii.

147

Food. The jackal is a predator-scavenger but, at the same time, it
readily eats all available carrion and refuse, and feeds on large quan-
tities of plant food, being, in some seasons, omnivorous.

In the Caucasus and Trans-Caucasus, it mainly hunts hares and
mouse-like rodents, as well as pheasants, francolines [F. francolinus|
(when they were numerous), ducks, coots [Fulica ата], moorhens
[Gallinula chloropus] and passerines. Birds suffer greatly from it at
the time of autumn migration and when wintering. It frequently attacks
domestic birds, including turkeys, more rarely lambs, sheep and goats
and one even attacked a newborn domestic buffalo calf (Dinnik, 1914;
Satunin, 1915). They willingly eat lizards, snakes, frogs, fish, molluscs,
insects, floating dead fish and various [kinds of] carrion. It frequently
feeds on fruits such as pears, hawthorn, dogwood, and “cones” of
Mespilus germanicus. It chooses sweet sorts and avoids sour. In gar-
dens, it destroys many grapes, eating watermelons, muskmelons and
nuts (Dzhanashvili, 1947). In 26 feces samples collected in summer,
1950, in the Ismaillinsk region (Azerbaidzhan), mammals were found
in 90.0% of the total sample, birds—27.0%, reptiles—23.1%,
amphibia—15.4%, insects—19.2%, plants—26.2% and carrion—7.7%
(Rukovskii, 1953). In 12 feces samples from Lake Shil’yan
(Azerbaidzhan) mouse-like rodents were found in 3.3% of the samples,
water vole [Arvicola}—25.0%, European hare [Lepus europaeus}
25.0%, carrion (wild boar, dog)—16.7%, coot—8.3%, discarded
fish—8.3%, insects (locust}—40.1% and squash seeds—16.7% (Pavlov.
1953).

In severe winters, when water bodies freeze, jackals cause great
losses among nutria, coots and ducks. In the winter of 1948/49 in
Trans-Caucasus, nutria occupied first place in the jackal diet (Table 4).

In such winters, mouse-like rodents, small birds and other foods
play a small role in the jackal ration. In winter, in the presence of
abundant prey the jackal kills more animals than it can eat, and caches
excess food. Jackals feed in a similar manner in Middle Asia and
Kazakhstan (Table 5). Rodents occupy the principal place and then
birds (20.0%).

Some role is also played by fish and arthropods, of which locusts
and darkling beetles are more frequently eaten (Chernyshov, 1954). In
some instances the stomachs of jackals are entirely filled with locusts
(Flerov, 1935).

In the nutritional ration of the jackal, wild and cultivated plants are
of serious significance. Thus, the stomach contents of 23 jackals caught

148

Table 4. Winter foods of jackal in Trans-Caucasus in the severe winter of
1948/49 (in % of the total number of samples investigated) (Pavlov, 1953)

Food designation Gagrsk region Kurdamirsk region
Abkhazsk ASSR Azerbaidzhan SSR
Before freezing During freezing period
30 samples 104 samples 17 samples
Nutria 26.6 89.4 47.0
Water vole 3:9 0.9 11.6
Brown rat 13:3 2.8 556)
[Rattus norvegicus |
Voles — — 5.9
Asia Minor gerbil — — 59
[Meriones tristrami]
Small mouse-like rodents 26.6 8:2 —
Birds 36.6 76.9 64.7
Including:
coot — —- 29.3
mallard — — 23.5
grebe — — 5:9
passerines — — 5.9
domestic fowl 10.0 — —
Fish 6.6 — —
Insects (beetles) — 1.9 5.9

Carrion of domestic animals 20.0 —

in October—January in tugais on the Vakhsh consisted completely, or
in large part, of the fruit stones of wild stony olive (Elaeagnus
angustifolia) (Chernyshev, 1948). In spring, jackals dig out and eat
plant bulbs (Flerov, 1935; Chernyshev, 1954), and later, roots of wild
sugar cane (Table 6).

Jackals living near the edge of the Karakum venture out into sandy
areas to hunt, where they catch gerbils, lizards and snakes. In the
period of drying of canals and irrigation ditches, they feed on fish and
frogs, and hunt muskrats. In tugais and gardens, jackals eat fruits of
wild olive (dzhida)*, mulberry and dried apricots, as well as water-
melons, muskmelons, tomatoes and grapes, to which they cause
appreciable harm.

In the lower Amu-Dar’ya, in the stomachs and feces of jackals,
were found remains of tolai hare, complex-toothed rat [Nesokia indica},

*Dzhida is a local name for wild (stony) olive—Sci. Ed.

149

114 Table 5. List of jackal foods in southwestern Tadzhikistan based on analysis

115

of gastro-intestinal tract samples (60 specimens) and feces (175 specimens)
(Chernyshev, 1958)

Type of food % of 235|| Type of food % of 235
samples samples

Animal food Reptiles and amphibians 2.8
Mammals Reptiles unidentified 0.8
Mammals, unidentified Lake frog 2.0
Carnivores Fish 6.8
Badger Fish, unidentified 2.0
Marbled polecat Gambusia 3.6

[Vormela peregusna] Common carp he
Rodents Arthropods 14.4
Rodents, unidentified Insects (majority are
Tolai hare [Lepus tolai] beetles, and others) 9.6
Field rat* Plant food 32.6
Turkestan rat Melon culture (musk-

[Rattus turkestanicus | melon, watermelon) 6.4
Red-tailed gerbil Wild olive fruits 14.4

[Meriones erythrourus] Wild cane sugar 4.0
House mouse [Mus musculus] (root, stems)
Mole-vole [Ellobius sp.] Bulbiverous plants 4.4
Birds Nightshade fruit 1.6
Birds, unidentified Wolfberry fruit 122
Pheasant Wheat 0.8
Duck Carrion 10.4
Coot

Passerines, unidentified

*Species not clear—Sci. Ed.

gerbils, pheasant and other birds, insects (locust and beetles) and rarely
plants (Salikhbaev, 1950). Where there are muskrats and nutrias, the
jackal destroys a great number of these rodents (Pokrovskii, 1953;
Kostin, 1956).

They visit the sea shore where they pick up the dead fish, wounded
waterfowl, molluscs, and feed on parts of marine animals discarded by
processors. They also feed near slaughterhouses, dumps, and cattle
burial places. In southern Dagestan in the 20’s, jackals regularly fed
near the railway lines, picking up food remains thrown out from pass-
ing trains by passengers. They were so accustomed to finding edibles,
wrapped in paper, that hunters placed poisonous baits along the railway
lines, wrapping it in newspaper (Heptner and Formozov, 1941).

114

115

150

Table 6. Seasonal variation in the list of principal foods of the jackal in
southwestern Tadzhikistan (Chernyshev, 1958)
Name of food Season of year

Winter Spring Summer Autumn

XII, I, Il Ш-У VI-VIII IX-XI*
Rodents 3! 4 3 3
Birds 3 4 4 3
Reptiles and amphibians 1 2 3 2
Fish 1 2 3 3

Arthropods

(majority are insects) 1 3 4 3
Wild plants 3 3 3 4
Cultivated plants 3 3 3 4
Carrion, refuse 3 3 3 3}

*Designates months of the year—Sci. Ed.
'Frequency of occurrence of food: very frequent—4; frequent—3; rare—2;
very гаге—1.

In summer the jackal, apparently, needs to drink regularly, since at
that time they always stay near water and are not found in waterless
deserts. In southeastern Turkmeniya, when considerable sections of the
Kushka [river] dry up, jackals, together with wolves and foxes, dig
holes in the bottom of the dry channel and drink the ground water
collected in them (Yu.K. Gorelov). Eating of watermelons, muskmelons
and grapes 1s also linked with the slaking of thirst.

Home range. In southwest Tadzhikistan, daily movements of jack-
als sometimes comprise 8—12 km, 1.e. the home range of one family in
tugai has a radius of up to 12 km. In Tigrovaya Balka Preserve, in an
area of 6000 hectares, from 10 to 23 jackal families were counted in
different years, 1.e., one family occupies an average of from 260 to 600
hectares (Chernyshev, 1954). In Georgia, at Lake Inkit, in the severe
winter of 1948/49, seven families of this carnivore occupied an area of
3 x 3 km (М. Pavlov, 1953). The average area for one family there was
130 hectares.

In case of food shortages in some winters, and also after large fires
in tugais and reeds, jackals in search of prey were obliged to undertake
movements of 40—50 km, regularly appearing in cultivated lands and
villages.**

**In Russian original, kishlak, a Turkic word for village—Sci. Ed.

116

151

Burrows and shelters. In the Caucasus and Trans-Caucasus, the
female usually gives birth in a burrow it has dug together with the male,
or she occupies the abandoned dwellings of fox and badger. The burrow
is dug some days before delivery, and the female and male take turns
digging. It is located in thick shrubs, on the slope of a ledge or gulley,
or on a flat surface. The burrow is simple in structure, since it possesses
one opening which runs to a depth at a small angle. The length of the
burrow is about 2 m, while the nest chamber is located at a depth of 1.0—
1.4 т. In Dagestan and Azerbaidzhan, jackal litters also were repeatedly
found in the hollows of large fallen trees (Satunin, 1915; Dzhanashvili,
1947). In Azerbaidzhan, lairs with pups were found under the roots of
trees and under large stones on river banks (Dzhanashvili, 1947).

The litter remains in the burrow for 2—2.5 months after which the
female leaves the lair with the young and begins to lead a roving way
of life. Regardless of the time of reproduction, the lair is built in dense
thickets of spiny bushes (blackberry, Jerusalem, or Christ thorn) or
reeds, and rarely in caves, crevices in rocks, hollows, old burrows,
water channels and other secluded places.

In Middle Asia, jackals construct lairs in the dense thickets of
tugais, and burrows are not dug (Chernyshev, 1954), but they have
been described for the Vakhsh tugais (Flerov, 1935). These burrows
usually have short, straight passages, dug under the roots of turanga or
wild olive, or directly in dense thickets of tamarisk and Nitraria sp.
One of these burrows, with a length of three meters or more, went
obliquely downwards for some distance and widened into the lair itself.
In front of the entrance there was a quite large heap of earth which was
greatly trampled down in the middle. At a distance of 140—200 т two
other holes, similar to the first one, were found. From all the holes,
well-packed paths extended through the tugai, which led to a common
network of trails. Surrounding the holes were many feces, partially
covered with earth.

In the tugais and cultivated lands of Tadzhikistan, jackals construct
lairs in the turf of a gigantic grass—plume grass [Erianthus], less
frequently in Nitraria sp., shrubs and in openings in reeds. There,
where vegetation may have been destroyed, jackals live in holes. The
female, male and young live in one lair (Chernyshev, 1954).

Daily activity and behavior. The jackal basically follows a cre-
puscular and nocturnal way of life, leaving its diurnal shelter before
sunset and returns with the light or even later in the morning. They also
often hunt by day. One may judge whether it is leaving to hunt by the

152

character of its howl. When one animal begins to howl, the others soon
answer. Howling is, apparently, also connected in part with the weather.
‚ Cases have been noted, when, in case of change in weather, jackals
break out in a prolonged and continuous howl. Jackals accompanied
with howls the ringing of church bells; the young start to howl when
they hear the sound of a siren, or whistle of steamship or train engine.
They usually howl in the evening and at dawn, but sometimes during
the day, even at midday.

The jackal is a daring and brave animal, and in places where it is
not pursued, less cautious and not afraid of people. It is easily caught
in traps or envenomed with poisoned bait. Many instances are known,
when, within sight of people, it attacked a domestic fowl, and regard-
less of the shouts and pursuit, takes it off. From field camps and from
houses it drags off various objects, which it thinks to be edible, such
as leather, boots, rags, venturing to steal them just a few steps from
sleeping people.

Jackals usually hunt singly, and less frequently in pairs and in [a
group of] 3—4 and only rarely do they form small packs. In the Trans-
Caucasus, at the end of summer and in autumn, packs of 8—12 jackals
were observed including, apparently, two or more families (Dzhanashvili,
1947). When embarking on a hunt, the jackal jogs along at a trot and
frequently stops to sniff and listen. On spotting prey, it stealthily con-
ceals itself, and quickly approaching, pounces and quickly makes a kill.
When hunting in pairs or packs, they run in parallel, overtaking the
prey together. For example, when hunting nutria or muskrats, they run
along both banks of narrow streams or channels, driving the frightened
prey from one to another. When lying in wait for its prey, it conceals
itself near a path or waterhole. In times of drought, they concentrate
near the drying lakes and channels and catch animals and birds on the
shore or in shallow water; they also catch fish there. During freeze-up,
they hunt waterfowl, nutria and muskrats coming out on the edge of the
ice or bank. They attack sleeping flocks of birds (ducks, little bustards
[Т. tetrax]) and especially many of them are destroyed in severe win-
ters when they are exhausted from starvation. In autumn, when the
canals and irrigation ditches are prepared for cleaning and are dry,
jackals together with foxes, jungle cats, badgers, crows, and magpies
successfully hunt fish and frogs that have collected in the remaining
pools of water, as well as muskrats forced to walk over significant
section of dry land. When water bodies freeze, in trying to catch
muskrats, they destroy their houses.

153

The jackals live in pairs, but are also found either singly, or in
pairs and families of 4—5 individuals.

Jackals, when taken young, quickly become tamed, following their
owner everywhere, and running to his call. They behave as an affec-
tionate dog would. Even adult animals can be tamed. In some places
the dogs do not disturb jackals and both animals could be observed
together on one garbage heap.

Seasonal migrations and transgressions. Regular seasonal migra-
tions are not characteristic of jackals. They usually wander only in
various directions searching for places with food in which they concen-
trate. Thus, in autumn they gather on the shore, where they pick up bird
remains. On sea coasts they are attracted by the carcasses of dolphins
or seals thrown out by processors. During fall, the number of jackals
in the tugai grows due to the arrival of animals from neighboring
cultivated lands. In winter, they wander into human settlements or
intrude deep into the desert. During such wanderings, they cross 50—
100 kilometers or even more.

While not truly regular migration, at the same time jackals often
settle at great distances from their usual range, and appear in regions
in which they were only a long time ago, or where they are completely
unknown (see above). In 1920—1922, when many cattle died of starva-
tion, jackals appeared in considerable quantity in western Kazakhstan,
especially in the valley of Ural river; in the north they reached nearly
to Orenburg (Sludskii, 1939). The jackal may appear around Orenburg
from the delta of the Amu-Dar’ya, passing along the shore of the Aral
Sea, through Ustyurt to the Emba and Ural; this way constitutes, in a
straight line, about 1000 km.

In the 30’s of the present century, the jackal was rare in the middle
course of Syr-Dar’ya (Aris’ region), but in 1942—1945, it became
common there, reached eastward to Chimkent, appeared in the Boroldsisk
mountains in the upper Bugun 150 km north of Chimkent (Sludskii,
1953). It remained in the latter region until the 50’s.

In the winter of 1948—1949, in Kustanaisk district, after an ex-
tended continuous snow cover and massive death of cattle on winter
pastures, jackals appeared in the summer of 1949, in Dzhangil’dinsk
and Amangel’dinsk regions of this district, on the Turgai river and at
Lake Sarykopa, and became common. Through 1952, their skins were
received in trade. The appearance of jackals in 1949 in the Turgais
steppes was accompanied by a massive inflight to this area of brown
[Gyps fulvus] and black [Aegypius monachus] vultures which were

154

also attracted by the abundance of carrion. On the Turgai river, jackals
could pass from the delta of the Amu-Dar’ya along the western shore
of the Aral Sea and the Bolshoi Barsuki sands, or also from the Syr-
Dar’ya, below Aris’, across the lower course of this river and the
Priaral’sk Karakum sands. In the first event they might have gone
about 1050 km, and in the second—about 1125 km.

In the severe winter of 1950/51 in southern Kazakhstan, jackals
appeared at the northwestern edge of the Betpak-Dala desert, in the
Dzhetykonur sands, having penetrated there from the Syr-Dar’ya up-
wards along the Sary-su river, and they remained there in limited
numbers until 1954; several animals, in that winter, succeeded in reach-
ing even the upper Sharubai-Nury river 40—50 km southwest of
Karaganda. The route thence from the Syr-Dar’ya is about 800 km. In
the lower Chu river, the jackal was encountered at the end of the past
century, then disappeared and again appeared in 1917. Later, individual
animals were caught there in 1927, 1936 and 1948.

In October 1924, a female of a pair was caught 60 km from
Tambov (Ognev, 1931). In order to reach Tambov from the lower
Terek along the Caspian Sea coast and upwards along the Volga and
its tributaries, jackals must have gone about 900 km. The appearance
of carnivores in new regions usually coincides with the mass death of
ungulates from starvation. It is not clear how jackals and vultures find
separate regions where there is much carrion.

In Bulgaria, jackals also sometimes migrate to the north and northwest,
appearing near Sofia, at the city of Kotel in Asenovgradsk region, at the
city of Dimitrova, and also intruding into Romania, Yugoslavia and Hun-
gary (Atanasov, 1953). In the winter of 1952/53 they appeared in Slovenia,
following, apparently, the herded sheep. Prior to this winter, jackals in
Yugoslavia were only found in Dalmatia and Macedonia (Berlikh, 1955).

Reproduction. Jackals live in pairs, united apparently, for life,
until one of the partners dies. The male helps in digging the burrow and
raising the young. In southwestern Tadzhikistan, out of 82 jackals, 39
were males and 43, females (Chernyshev, 1958).

In the Trans-Caucasus, estrus begins in the first half of February,
and in warm winters, at the end of January. The mating period conti-
nues for 26—28 days (Dinnik, 1914; Satunin, 1915; Dzhanashvili, 1947).
In Tadzhikistan and Uzbekistan, the rut manifested itself at the same
time also, and extended to the first days of March (Chernyshev, 1954;
Kashkarov, 1931; Salikhbaev, 1950). In Bulgaria, it extends from the
middle of January to mid-February (Atanasov, 1953).

118

SS)

Gametes in adult males and females do not mature at the same
time. Active spermatogenesis in males appears 10—12 days earlier than
estrus in females. During the mating period, testes weight increases
three times.* In Tadzhikistan, average weight of testes in January
was 6.1 ©, in February—13.3, in March—13.0, in April—7.7 and in
August—4.4. In a male, caught on 6 March 1948, with noticeable
degradation of the testes, sperms were still detected. Ovaries increase
insignificantly in the estrus period (from 0.7 g in January to 0.8—0.9 ©
in February—March) (Chernyshev, 1954). Estrus continues 3—4 days. If
the female is not mounted during this period, her receptive condition
will be interrupted for 6-8 days, after which estrus returns and again
continues 3—4 days. If the female does not become pregnant Бу that
time, estrus will not recommence until the next year. Mating can be
observed at any time during the diurnal cycle and is accompanied by
an attachment [mating tie] which lasts for 20-45 minutes (Dzhanashvili,
1947).

Jackals howl particularly often during the mating period. The fe-
male coming into heat for the first time is pursued by several males,
and quarrels develop among them. The duration of pregnancy is 60—63
days (Bogdanov, 1873, Pocock, 1941, Brink, 1957); according to
observations at Tbilisi 700—62—63 days (Dzhanashvili, 1947). In
Tadzhikistan a considerable number of barren females (5 out of 15)
was noticed in different years (1948, 1950). In all of these barren
females, helminths were found in great quantities (Chernyshev, 1954).

In the Trans-Caucasus, birth of the young usually takes place from
the end of March to the last days of April. The earliest parturition was
noted on 28 March 1937 and the latest, on 25 April 1931 (Dzhanashvili,
1947), but sometimes females deliver in the first half of May (Satunin,
1915). In Tadzhikistan, Uzbekistan and Bulgaria parturition occurs in
April and the beginning of May. Only once, a female gave birth to 8
pups on June 11th (Chernyshev, 1958; Atanasov, 1953). Only one litter
per year is produced.

In the Trans-Caucasus, the number of young in the litter ranges
from 3 to 8 (Dinnik, 1914; Satunin, 1915; Markov, 1934; Dzhanashvili,
1947). Satunin (1915) dug out several burrows, in one finding 5 pups,
and in the remainder, 4. According to observations in nature and in
zoos, the litter usually contains 5 pups, rarely 3—4 or 6-8. In
Tadzhikistan, there are 3—7 (average 4.8 pups; Chernyshev, 1958). In

*The weights given in the next sentence suggest that this should read “two итез”—
Sci. Ed.

119

156

Uzbekistan, there are 2—8 pups in a litter (Kashkarov, 1931; Salikhbaev,
1939, 1950). In the majority of regions in Bulgaria, there are 4—7 pups,
but in Michurinsk region, only 3—5 (Atanasov, 1953). In India, a litter
contains an average of 4 pups (Blanford, 1888).

The female nurses the young for 50—70 days (Caucasus;
Dzhanashvili, 1947), in Tadzhikistan up to 90 days. The lactation
period in some females ends in the middle of July and in others, in the
first days of August (Chernyshev, 1958). The young begin to eat meat
starting at the age of 15—20 days. Only in rare instances does the
mother feed the pups regurgitated food. At the end of the lactation
period, the female drives the young away from her by growling and
biting. The latest-born remain with the female until the first half of
autumn, after which they leave her and start leading an independent
life, either singly or in groups of 2—4 individuals.

Sexual maturity, based on observations in the Tbilisi zoo, takes
place in females at an age of 10—11 months, but in males at the age
of 21—22 months. In males 10—11 months old, the testes are undevel-
oped and contain no sperm (Dzhanashvili, 1947). In Tadzhikistan, males
are said to become sexually mature at the age of 10—11 months
(Chernyshev), a statement which is not true. Duration of life is 12—14
years (Brink, 1957).

Growth, development, and molt. Pups are born with a pelage of
soft fur, the color of which varies from light-gray to dark brown. At the
age of one month, the juvenile pelage is replaced. At first, guard hairs
grow on the head, back and rump, then on the anterior and posterior
limbs and, finally, on the tail. Fur color of the young changes, becom-
ing reddish with black speckles. Young caught by the end of August
and the beginning of September hardly differ from adults in fur color.
By October and November, the young already have full winter fur
(Chernyshev, 1958).

Young are born with closed eyelids. In the Tbilisi zoo, ш 39 pups
(78%) the eyelids opened on the 9th and in the remainder—on the 10th—
17th day. Eye-opening is delayed in weak and sick pups (Dzhanashvih,
1947). In Tadzhikistan, based on observations of 9 pups, eyelids opened
on the 8th—1 1th day (Chernyshev, 1958). Ears in healthy pups open on
the 10th—13th day and become erected only on the 25th—30th day.

The last [3rd] and middle [2nd] deciduous incisors in the upper jaw
first erupt on the 11th—14th day. The inner [1st] deciduous incisors of the
lower jaw are the last to erupt, on the 17th—19th day. At that time, the
deciduous canines appear. The replacement of the deciduous by permanent

157

incisors is completed at about the age of 4 months, and the third and final
molar of the lower jaw erupts at an age of about 5 months. The times of
eruption of deciduous teeth and the order of their replacement by the
permanent ones is related to lactation, protracted feeding of the pups, and
the time of their transition to an independent life.

Young jackals grow quickly. At the age of two days their weight
is 201-214 g, at one month, from 560—726 g, and at 4 months, 2700—
3250 g. Young caught at the beginning of September weighed 4200—4370
g, and by the beginning of October, their weight reaches 3/4 of adult
weight (Chernyshev, 1958).

The jackal molts twice yearly—in spring and in autumn. In the
Trans-Caucasus and Tadzhikistan, the spring molt begins in the middle
to end of February, in warm winters, in the middle of that month; in
cold winters, it starts in mid-March and ends at the beginning to middle
of May. In healthy animals, pelage replacement extends for 60—65 days
in spring. Weak and sick animals do not succeed in completing molt
during the entire summer; at the beginning of winter, half of the coat
consists of spring—summer hair and half is winter fur (Dzhanashvili,
1947).

Spring molt starts with the head, the anterior and posterior limbs,
then spreads to the sides, back, chest, belly and rump, and the last to
molt is the tail. After spring molt, only scattered coarse guard hairs
remain on the skin and underfur is absent. Autumn molt occurs from
the middle of September. Shedding of summer pelage and growth of
winter guard hairs and underfur take place simultaneously. Autumn
molt starts with the rump and tail and spreads to the back, sides, belly,
chest, limbs and head. Full winter fur has grown out by the end of
November (Chernyshev, 1958). In the Caucasus, the skin is prime from
November to March (Vereshchagin, 1947).

Enemies, diseases, parasites, mortality, and competitors. Earlier,
enemies of the jackal everywhere were the tiger and leopard. At the
present time, the first is extinct in the jackal’s range, and the second
is extremely rare. The leopard hunted jackals especially frequently. At
the same time, jackals regularly fed on the remains of these cat’s prey,
and once, in Tigrovaya Balka, they ate a dead tiger. The wolf also
should apparently be considered an enemy.

Its serious competitors everywhere are the jungle cat, red fox, wolf,
and in the Trans-Caucasus, the forest wildcat [Felis sylvestris], and to
a lesser degree, the recently acclimatized raccoon [Procyon lotor] as
well. In Middle Asia and Kazakhstan, the steppe wild cat [Felis libyca]

120

158

is also a competitor. The small number of red foxes occurring in tugais
may explain the abundance of the jackal there (Chernyshev, 1958). At
the same time, in deserts and foothills, red fox and wolf apparently oust
the jackal. Usually, where there are many wolves, jackals are absent,
and vice versa.

Among the infectious diseases of jackal, carnivore distemper and
rabies are established (Sludskii, 1954; Chernyshev, 1958). Epizootics
of carnivore distemper in the winter of 1948-49 in Tadzhikistan
(Tigrovaya Balka) affected not only jackals, but was also observed
among shepherd and hunting dogs, the majority of which died. In south-
western Tadzhikistan, of 9 jackals examined, leishmania was found in
liver and spleen smears of three, and was supposedly identified as
Donovan’s leishmania, which induces visceral leishmaniasis in humans.
Jackals infected with leishmania suffered little from this infection
(Latyshev and others, 1947).

Jackals are heavily infested with helminths. In southwestern
Tadzhikistan, 15* species of cestodes, roundworms and acanthocepha-
lans were discovered: Sparganum mansoni, larval form of
Diphyllobothrium mansoni, Taenia hydatigena, Taenia pisiformis,
Taenia ovis, Hydatigera taeniaeformis, Diphylidium caninum,
Mesocestoides_ lineatus, Ancylostoma caninum, Uncinaria
stenocephala, Dioctophyme renale, Toxocara canis, Toxascaris leonina,
Dracunculus medinensis, Filariata gen. sp., [and] Macracanthorhynchus
catulinum. Jackals infect water bodies with the parasitic worm larvae
(D. medinensis) and may be responsible for dracunculosis in humans
(Chernyshev, 1954). Moreover, they play some role in the spread of
coenurosis of sheep and large horned cattle. In Uzbekistan, 10 species
of helminths were found in the jackal, of which, in addition to 7 species
mentioned for animals in Tadzhikistan (No. 6-12, see above), were
found: Spirocerca lupi, Rictularia affinis and R. cahirensis (Irgashev,
1958).

Jackals in Tadzhikistan are infested with ticks in April—September,
while in November—February, these parasites are only found in few
individuals and in small numbers. The following species of ticks were
found: Ixodes sp., Rhipicephalus turanicus, R. leporis, R. rossicus, R.
sanguineus, R. pumilio, R. schulzei, Rhipicephalus sp., Hyalomma
anatolicum, H. scupense, H. asiaticum, Hyalomma sp. Most frequently,
R. sanguineus parasitizes the jackal in great quantities. The role of
jackals in nourishing and spreading ticks is great. Four flea species are

*16 are listed—Sci. Ed.

159

Fig. 43. Track of jackal on silt. “Tigrovaya Balka”, preserve at Pyandzh,
Tadzhikistan. 15 October 1960. Photograph by A.A. Sludskii.

known on jackals: Pulex irritans, Xenopsylla nesokiae (a flea of the
bandicoot rat), Ctenocephalides canis, С. felis and one species of
louse—Trichodectes canis (Chernyshev, 1954).

In severe deep-snow winters, jackals often die from emaciation and
exposure. In Azerbaidzhan, in severe winters, after the large lakes are
covered with ice, jackals penetrate far into the dense growths of reeds
and cattails and stay in them during the entire frozen period, not going
to shore. At times of sudden thaws, they are stranded on the ice and
die. Thus, in December 1946, when a thaw commenced, the animals
which were living in the reeds were isolated from the land by water and
drowned (Vereshchagin and Dyunin, 1949). In the severe winter of
1948-49 at the Shil’yansk lakes, as a result of sudden thawing, the
animals became isolated from the shore and drowned at the borders of
the impassable flooded reeds in their attempts to reach dry land. Other
cases were observed when jackals, failing to reach the shore, remained
starving for a long time on intact sections of ice (Pavlov, 1953). Near
the Caspian Sea jackals sometimes die in great numbers in autumn and
winter moryan* which quickly fill coastal thickets, and in times of
floods. Possibly, they also suffer from fires in the turgais and reeds.

*Violent on-shore storm — Sci. Ed.

121

160

Population dynamics. These are little studied. Judging by the data
of fur buyers, the number of jackals is subject to considerable fluctua-
tion. Thus, in Turkmeniya, fur purchases varied by nearly 3—4 times
during the period from 1950 to 1956 (1400—5500 items). They fell
especially greatly in 1954—to 1430—following the severe winter of
1953—54. Fur purchases considerably increased in 1956, reaching 5427
skins, following 1955, a year which was favorable for reproduction
because of abundant food.

In Tadzhikistan, minimal fur purchases during the ten years was in
1951—258 pieces after the severe winter of 1950—51, and the maxi-
mum, as in Turkmeniya in 1956—1403 skins. In Tadzhikistan, fur
purchase, and consequently the number of jackals, fluctuated approxi-
mately by six times. In the lower Amu-Dar’ya in the decades of the
30’s—50’s the catch of jackal skins varied by 2 times and was also
related to the degree of severity of winter. In the south of Kazakhstan,
jackals were relatively common in the 20’s and 40’s of the present
century, during periods when there was much carrion in the desert.

In 1926-1929 in Kazakhstan, from 100 to 250 jackal skins were
obtained; at the beginning of the 30’s, this number dropped to 25—30
pieces; after which a few individuals were caught; but in 1942—1945
the number of jackals started to increase. By 1947, jackals were com-
mon in the most southerly regions. In this year, 104 jackal skins were
tanned in Kazakhstan. In the following years, their number again
dropped. However, these changes were mainly connected with their
migrations from the south.

Field characteristics. The jackal is half the size of the desert wolf
and nearly twice as large as the desert red fox, and is high in the limbs.
Ears are erect, but more blunt than those of foxes. Tail short and
1/3 the body length.

Feces are sausage-like, and are found in noticeable piles, as the animal
leaves them in one and the same place for several times. It often howls. The

“howling is similar to a wail, and can be rendered by the sounds “Ai-yai!

Ai-yai!”—high, whimpering. It is similar to that of dogs, but is higher and
more varied and is differentiated by a “plaintive” intonation. To the howl
of one jackal, those nearby answer (A.S.).

Practical Significance

The jackal is a fur-bearer, but its fur is of low value, characterized by
sparse, coarse pelage and has a monotonous unbeautiful color. Already

122

161

in the last century the jackal was utilized (Silant’ef, 1898). In the
Trans-Caucasus, in the Zakatal area, 300 jackals were captured in
1896, and in the B[olshoi] Trans-Caspian district, the jackal was hunted
in many jurisdictions, and in Mervsk, in the eighties, about 200 animals
were caught yearly. The total taken in Russia in that period was more
than ten thousand jackals. Jackal skins were sent to the Nizhegorod fair
(they were not sent to other fairs) in the following numbers:

Year 1880 1881 1883 1884 1885 1886 1887
Number of skins (in units) 8000 10,000 3000 4000 4000 3000 2800
Year 1888 1889 1890 1891 1892 1893
Number of skins (in units) 3000 3000 3500 4000 4000 3000

In the decade of the 20’s in this century, the following numbers of
skins of this animal were tanned in Trans-Caucasus (Markov, 1934):

Years 1924— 1925— 1926— 1927— 1928— 1929- 1931 1932
1925-1926 927 1928-1929 +1980

Number of skins 4386 6159 4027 11793 9334 13635 11819 13384

In Middle Asia and Kazakhstan lesser amounts of skins were pre-
pared (Tables 7, 8).

In the entire Soviet Union, at the beginning of the 30’s of the
current century, about 20—25 thousand were tanned yearly (Naumov
and Lavrov, 1941; Kaplin, 1960). However, the stocks of this animal
were significantly underutilized. In 1940, 10.4 thousand furs were pro-
cessed. At present, it is possible to increase the size of the jackal take
by a minimum of three-fold. The world production of jackal skins in
1907—1910 was 20-25 thousand items (Asia 5.0, Europe 5.0, Africa
10.0; Kaplin, 1960). Previously, the majority of skins tanned in the
USSR were sent abroad, mainly to the USA. Since 1949, the skins have
been utilized inside this country.

The skins are not graded according to a fur standard, though those
obtained from the Transcaucasus possess coarser fur of reddish-brown
color, while those from Middle Asia differ in their softer paler fur.
They are divided into four sorts, and used in the manufacture of cheap
collars, women’s coats and dokhas [fur-coats]. They are usually dyed.
Suede is prepared from the skins of grade IV.

A special harvest of jackals does not exist. The animal is caught
incidentally to the hunting of other animals, by means of traps or by

162

ves v9L c0rl LS8 088 816 9LS

8ST == Ее Е USS WyzpeL
895: У89Р v89Pr 6009 ССР C9SP 965? 819Р 1919 a = 455 4992
SOLC 156 [Ст 8105 0571 Legl SOSI SL81 $907 6861 9SLT USS vouryin yp
solqnday
8561 1561 9561 $561 7561 5561 561 1561 0561 6761 8761 uoruf)

ав.
a a es

(SHUN Ut) 8561-8761 рома ay) UF ау a[PpIA ит saseYydsind Uys [eyoEL "8 Е

мФ— eee

111 РЯ 8rz == — 415$ yyezey
€ I С = = USS 218 ИУ
повецчоит ON Uss usWyIN |

9IPZ LL9Z €rSZ — 8151 USS Ayzpey рие yoqzq

Е Е ee
6061-8061 8061-1061 1561-9061 9561-5061 — 6061-2061 Sinaia
Е ee eS eee

SIRO A uoluQ

a a ee eee

(sjiun ul) UBIsYyYeZey рие в15у э|ррид! Ul saseysand шуб ЕЛЕ */ ЭЧЕТ

123

163

shooting while on drives. When traps are employed, they are set near
carrion, on garbage dumps (in such places the beast pays little attention
to the odor of metal) and in shallows along shores; as a bait, a large
live fish is tethered on a kukan* (string).

In the Trans-Caucasus, jackals are hunted with large fishing hooks
which are suspended from a metallic cable or wire at a height of 75—
100 cm from the ground, in such a manner that the predator can reach
it only by jumping. The hook 1$ baited with a piece of carrion. On
seizing the bait, the animal is hooked by the lip or jaw (Markov, 1934).
The jackal is snared by wire nooses placed on their trail. Poisons are
also utilized (see above concerning food).

The jackal is a serious pest of the agricultural and hunting econo-
mies. In the Trans-Caucasus, “jackals must be considered the most
harmful destroyers of game” (Dinnik, 1914), causing the most severe
losses to the hunting economy, not excepting the wolf (Markov, 1934).
They destroy, in large quantities, newborn roe deer, hares, and nutria
as well as pheasants, francolins, gray partridges, little bustards and
great bustards during overwintering, and waterfowl (Satunin, 1915; M.
Pavlov, 1953; Katubadze, 1956; Burdzhanadze, 1959). The jackal is a
dangerous enemy of the nutria, completely destroying them in shallow
water bodies (M. Pavlov, 1953). They are very harmful to the muskrat
trade. In the Amu-Dar’ya delta, during the winter of 1948—49, in the
feces of this predator the remains of muskrat constituted 12.3% of all
examined samples. In particular places, jackals destroyed up to 71% of
the muskrat houses, of which about 16% froze and became unsuitable
for occupation (Pokrovskii, 1953). It causes also great harm to the
muskrat trade in the Khorezma district where muskrat houses are easily
accessible to it. The jackal also damages the muskrat harvest by eating
muskrats caught in traps, and skins hung up for drying (Kostin, 1956).

The jackal is very harmful to poultry and carries off lambs and kids
(Satunin, 1915). It eats ripe grapes in such quantities that earlier, in the
Bol. Lenkoran vineyards, owners removed the flowering branches hang-
ing near the ground because the fruits will in any case be eaten by
jackals (Satunin, 1915). The jackal destroys watermelon and melon. In
one melon field in Tadzhikistan, the jackals destroyed, ate and spoiled
70 ripe melons in one night (Chernyshev, 1958). In the Trans-Caucasus
and Middle Asia, the jackal, as much as the red fox and wolf, serves
as a carrier of rabies in nature.

*Local word?—Sci. Ed.

124

164

Cases are known in which rabid jackals have run into villages,
where they have bitten man and domestic animals (Bikhner, 1905). The
value of jackals as eaters of small rodents and insects is inconsiderable.

The jackal may be killed the year round, until such time as their
numbers proceed down to a minimum. In all republics, there exist
decrees permitting their hunting throughout the entire year. (A.S.)

1758.

1792:

1804.

1831.

1847.

1863.

1875.

1882.

1907.

WOLF
Canis lupus Linnaeus, 1758

Canis lupus. Linnaeus. Syst. Nat., Ed. X, 1, p. 39. Upsala,
Sweden.

Canis lupus albus. Kerr., Animal Kingdom, p. 137. Lower
Yenisel.

Canis lupus communis. Dwigubski. Prodromus faunae Ross., p. 10.
“Russia” —apparently, the northern forested section of European
Russia (the steppe wolf Dwigubski inserts separately; see below).
Canis lupus campestris. Dwigubski. Ibidem, p. 10. Cis-Cauca-
sian steppes.’

Canis pallipes. Sykes. Proc. Zool. Soc. London, p. 101. Deccan,
India.

Lupus laniger. Hodgson. Calcutta Journ. Nat. Hist., 7, p. 474.
Tibet. Nec Canis laniger H. Smith 1840 (domestic dog of Puget
Sound Indians).

Canis chanco. Gray. Proc. Zool. Soc. London, p. 94. “Chinese
Tatary”. Tibet.’

Canis arundinaceus. Karelin. Tr. SPb Obshch. Estestvoisp., 6,
p. 81. South Ural steppe. Nomen nudum.

Canis lupus var. desertorum. Bogdanov. Ocherki prirody
Khivinskovo oazisa 1 pustyni Kyzylkumy, р. 30, Kyzylkum.
Lupus filchneri. Matschie. Wiss. Ergebn. Exped. Filchner nach
China u. Tibet, 10, 1, p. 153. Si-ning to the east from Kukunor.
Lupus karanorensis, Matshie. Ibidem, p. 157. Khala-chu in Gobi
(a little west of Sa-chzhou. V.H.).

Lupus tschiliensis. Matschie. Ibidem, р. 160. Shanghaiguan’ on
shore of Lyaodunsk gulf.

Рог reasons, see later in section “Geographic Variation”.

1922.

1923.

1923:

1926.

1929.

1930.

ТОЭ:

1935:

1936.

LOST:

165

Lupus altaicus. Noak. Zool. Anz., 35, p. 465. Teletskoe lake
and Chulyshmansk mountains, Altai.

Canis lupus var. orientalis. Auct. Dybovski. Archiv Tovar,
Naukow. Lwowe, 1, p. 350. Forest of Eastern Siberia. Nec
orientalis Wagner 1841. Nomen nudum.

Canis lupus var. argunensis. Dybowski. Ibidem, р. 350. Steppes
of Dauriya. Nomen nudum.

Canis lupus var. kamtschaticus. Dybowski. Ibidem, p. 350.
Kamchatka. Nomen nudem.

Canis lupus cubanensis*. Ognev. Biologicheskie izvestiya, 1, p.
114. Mountains of Maikop area, Western Caucasus (district of
present Caucasian preserve, at sources of Kuban tributaries—
Laba and Belaya).

Canis lupus turuchanensis. Ognev. Ibidem, р. 115. Turukhan
territory (right bank of lower Yenisei).

Canis lupus dybowskii. Domaniewski. Annal. Zool. Musei
Polonici Hist. Nat., 5, p. 52. Kolygino, southwestern coast of
Kamchatka.

Canis lupus bactrianus Laptev. Opredelitel’ mlekopitayushchikh
Srednei Azii, 1, р. 51. Surkhandar’insk region.

Canis lupus coreanus. Abe. Journ. Sc. Hiroshima Univers. Zool.,.
1, p. 33. Near Seoul, Korea.

Canis lupus hattai. Kishida. Lansania, 3, No. 25, p. 73. Hokkaido,
Japan.

Canis lupus rex. Pocock. Proc. Zool. Soc. London, p. 659.
Hokkaido, Japan.

Canis lupus dorogostaiskii. Skalon. Izv. Gos. protivochumn.
inst. Sibirii 1 Dal’nevo Vostoka, 4, р. 193, Borzya, Eastern
Trans-Baikaliya.

Canis lupus hajastanicus. Dahl. Dokl. Akad. nauk Armyansk.
SSR, 14, No. 3, р. 91. Vicinity of Erevan, Trans-Caucasus. (У.Н.).

Diagnosis

Body length of adults not less than 90 cm, tail length about half that
of body. Length of skull more than 200 mm. Arciform line formed by
anterior border of both nasals, has no process in middle (along line of
suture between nasal bones). At anterior inner sides of auditory bullae,

*In Russian original, misprinted as cubanenesis—Sci. Ed.

125

166

swellings in form of ridges found on basioccipital. Cingulum on outer
side of first upper molar narrow and weakly developed, with constric-
tion or gap in middle. (V.H.).

Description

The wolf is a slender, well-proportioned, powerful animal. It has a
strong trunk with sloping back, with high shoulders and a lower, but
strong and wide croup. The rib cage is large, deeply descending, the
abdomen is pulled in and the neck is powerful and muscular. The limbs
are long and strong, the paws are relatively small and the digits are
tightly compressed (“in a lump”).

Head large, heavy, with strong jaws, a long but not pointed, muzzle,
and broad forehead. Eyes are widely separated and small. Supraorbital
arches are well developed and therefore the eyes appear to be seated
deeper as if slightly squinting, even somewhat slanting. Ears relatively
small, triangular in form with pointed tip, directed forward and widely
separated, and therefore the wolf’s head looks especially “broad in the
forehead”. The animal usually carries it [the head] a bit hanging, not
higher than the level of the back, and appears slightly stooping, with
its high shoulders. Only the watchful wolf raises its head high.'°

Tail fairly large, fluffy, hanging down to the tarsal joint. It looks
as if broken at the base and in the standing and calmly moving wolf,
it hangs directly downwards. Only during fast galloping does the wolf
somewhat raise it, and carries it “outstretched” but not higher than the
back level. In the live animal, the tail moves little and looks very heavy
(in hunting language it is aptly called a “billet” [chunk of моо4]). In
known psychic states (glad excitement, affection) the wolf wags its tail,
although not entirely as the dog does. When in fear, it puts its tail
between its legs as does the dog. Claws black. Eyes yellow. Teats, 5
pairs.

The winter fur is very dense and fluffy with short underfur and
long, quite coarse guard hair. The tail is very densely covered with long
hairs, which are considerably shorter at the root than along the remain-
der of its length. In the south, the fur is scarce and coarser, in the
mid-zone, dense and luxuriant, but fairly coarse, and in the north it is
longer, denser, more luxuriant and softer.

"The wolf is an animal of high psychic [sic] level, and possesses rich mimicry.
Special study showed that the wolf has ten expressions of the “асе” corresponding to
specific psychic conditions.

126

Fig. 44. Wolves, Canis lupus L. Sketch by A.N. Komarov.

The longest hairs are found on the back, mainly on its anterior part,
and on the neck. On the shoulder there is usually a region of especially
long hairs, and on the upper part of the neck the long hairs form a kind
of crest. The anterior part of the head, including the forehead, is cov-
ered with short hairs, while on the remaining part, the hairs are longer.
On the cheeks, the hairs are elongated and form “tufts”—small side-
whiskers. The limbs, to elbows and a little above the calcaneal joint,
are covered by short, elastic, closely adjacent hairs. The ears are cov-
ered with short hairs and strongly project from fur. The summer fur in
all latitudes is much more sparse and shorter than that of winter, and
is coarse and harsh.

Coloration is monotone, in various parts of the country monotypic,
and geographically varies comparatively little. Individual variation is
fairly great, but affects details, while the general color tone and distri-
bution of colors are constant. Seasonal differences in coloration occur,
but are not sharp, and vary in different parts of the range. In some
areas, in connection with color fading in winter, the color somewhat
(sometimes considerably) changes (brightens). Among Soviet wolves,
melanists, albinos and erythristic are found, but are very rare. Their
appearance sometimes results from hybridization with domestic dogs.

168

Fig. 45. Head of adult male wolf. Krasnoyarsk territory. Photograph by
V.V. Kozlov.

The wolf of southeastern North America, С. /. niger, has diphasic
coloration—normal and black.

Sexual differences in color are absent. The coloration of the adult
Middle Russian wolf in summer fur is composed of a mixture of ocher-
ous and rusty ocherous tones with light gray. Over the entire skin,
especially on the surface of the back, there some admixtures of black,
as a result of the black ends of the guard hairs. The muzzle, approxi-
mately to the level of the eyes, is pale ocherous-gray, the circumference
of the lips and the lower parts of the cheeks are white. The space
between eyes, the forehead, top of head, occiput and the region under
the eyes and between eyes and ears are gray, only with a light reddish
film. Around the eyes there are small ocherous rusty fields (rings). The
backs of the ears are rusty-ocherous with a mixture of blackish-brown,
and their inner surfaces are covered by dirty white hairs. The chin and
throat are pure white. The neck is ocherous, somewhat overlaid with
black (dark ends of hair) on the upper side.

127

169

Long, dark (black) guard hairs along the spine form a well-defined
black pattern in the form of stripe which is especially bright and broad
in the anterior part of the back. A quite large amount of hairs with
black tips is also present on the shoulder blades, and in part on the
upper part of chest, and the sides on the posterior part of the body.
However, a dark frosting in the form of obvious “dorsal saddle ” is not
formed. The coloration of the sides of the body and the outer parts of
limbs is pale dirty ocherous and the inner sides of the limbs are white.
Belly and inguinal region are white with an ocherous frosting. On the
anterior side of the fore legs there is usually a sharply outlined and
well-defined longitudinal stripe.

Coloration of the tail corresponds to that of the sides of the body—
bright dirty ocherous. On the dorsal side, especially on the basal third,
there is a strong admixture of dark (black, black-brown) hairs; the tip of
the tail is often black. Hair length in the middle of the back is 60—70 mm.

The winter fur retains in general the main tones and also distribu-
tion of the variously colored parts. However, general coloration is
brighter as a whole in connection with the fact that the underfur is more
prominent; the tinge of black hairs on this background is more striking,
appearing as a sort of weak dorsal saddle; ocherous tones appear weaker
and the smoky-gray is more strongly developed. If in some individuals,
the ocherous tone is clear and quite intensively developed, in some
others it is weak and the animal looks very light and gray. The length
of the guard hairs on the shoulders does not usually exceed 90 mm, but
may reach 110—130 mm.

Age variation in characters of the fur and in coloration is in the
first year well expressed. Young wolves in their first pelage are cov-
ered with dense, short (in the middle of the back 20—30 mm), very soft
“downy” fur of dark-brown or grayish-brown color. This coloration is
quite evenly distributed on the entire body. The region around the eyes
and the inner surface of the body are somewhat lighter, while the
muzzle and the lips, on the contrary, are darker. On the belly the hairs
are dirty gray with brownish tints and on the chest, between the ante-
rior limbs, there is a lighter area. The color of the tail, which is covered
by short hairs, corresponds to that of the trunk. A white tip on the tail
never occurs. The claws are light.

This juvenile pelage quickly begins to change and at the end of
summer the young wolf, still far from attaining the size of the adult (not
less than two times smaller) has light dirty grayish-ocherous fur, coarse
and sparse. Reddish tone is not developed and black guard hairs are

170

Fig. 46. Adult male desert wolf, weight 31 kg. Betpak-Dala desert,
Kazakhstan. 24 May 1955. Photograph by A.A. Sludskii.

absent or few. This coloration is quite evenly distributed on the body
and the light or, on the contrary, dark areas are not differentiated. The
claws become black.

From this second pelage, the hairs of which grow greatly by
autumn, the young (“growing”)* wolf passes into the first winter
pelage. It corresponds to the winter fur of adult animals, but differs in
the greater monotony of the grayish dirty-ocherous color with less
development of black and reddish. The color of the wolf in the second
year (“yearling”)**, both in summer and especially in winter, does not
differ from that of older animals."

*The Russian word pribylnoi, while literally meaning “growing” or “increasing”,
is best considered the equivalent of “juvenile’—Sci. Ed.

**Russian, pereyarka—Sci. Ed.

''Numerous hunters have remarked on age differentiation of wolves on the basis
of color, as well as have the notes of old authors (Lorents, 1890; Sabaneev, 1877 and
others) but these are not reliable and invalid (Ognev, 1931; Heptner, 1947; and Heptner
and Turova, 1951).

128

ТИ

The skull of the wolf is characterized Бу massiveness and generally
large dimensions. This is the largest form in the family. The facial
portion, in connection with strongly developed teeth, is relatively long
and massive; the cranial part of the skull is comparatively small and
weakly swollen, and is considerably shorter than the facial part, and the
braincase is relatively small. The nasal bones are long, their posterior
ends extending to the level of the orbits. Each bone, at the front, is
notched in an arciform manner in such a way that a common protuber-
ance is not formed at the line of contact of both bones anteriorly (Fig.
15). Along the whole length of the line of contact of the nasal bones
with each other a longitudinal depression (groove) is present.

The premaxillae gives off a large dorso-posterior process which,
however, does not reach the frontal bones. The posterior borders of
nasals and maxillary bones lie approximately on the same level. The
zygomatic arches are massive and widely separated, especially poste-
riorly. The supraorbital processes are large and massive and strongly
protrude laterally. The facial portion is quite elevated anteriorly and in
the middle and posterior parts of the nasal bones the profile is some-
what concave and the forehead abruptly rises upwards, reaching its
greatest height in the region of the supraorbital processes.

The frontal region is wide, slightly concave in the middle and
convex on its margins. Behind the supraorbital processes, the cranium
is compressed. The sagittal crest 15 well developed; it bifurcates ante-
riorly, demarcating the frontal area from the sides and continues to the
border of the posterior part of supraorbital processes. The occipital
crest is strongly developed and overhangs the occipital region of the
skull. The bones of the auditory bulla are of moderate size, thick-
walled, and their anterior inner parts are not directed to each other, but
diverge laterally; in this part, at the base of the jugal, are found small
elongated ridges or swellings.

The tooth row is very powerful, carnassial teeth are massive and
the canines are strong, relatively short, but with a wide base.

Age variation of the skull is very great and is reflected mainly in
the development of the crests, the relative increase in the facial portion,
increased convexity of the frontal region, and the increased constriction
of the cranium behind the orbits.

The wolf cub skull still possessing deciduous teeth is characterized
by the following characters: facial region of the skull is very short and
much shorter than the cranial; width of the skull in the region of
carnassial teeth is great; zygomatic arches are situated very close and

130

172

are weak; cranial portion of the skull is relatively large and swollen;
crests are absent; contour of the braincase region is circular (no pro-
jections in the upper part of the occiput); no constrictions behind the
orbits; supraorbital processes not defined; frontal region rises slightly,
and projections in this part of the skull are absent; auditory bullae are
relatively large and more rounded; the coronary processes of the lower
jaw are sharply curved backwards; and the angular processes are small.

The skull of the juvenile wolf immediately after tooth replacement
(in the first autumn of life) has a skull that appears similar to that of
the adult animal, but differs in the following characters: nasal portion
is somewhat shorter; skull width in the region of carnassial teeth is
somewhat greater; zygomatic breadth is less; region of the braincase is
relatively a little longer; crests are absent, and only in the occipital
region is the posterior part of sagittal crest defined; supraorbital pro-
cesses are small, short and weakly pointed; constrictions behind the
supraorbital processes are less, and bones of the auditory bullae are
relatively larger.

The skull of the very old wolf, as compared to that of the adult, is
characterized by a relatively still more elongated facial portion, mas-
sive and very widely separated zygomatic arches, high and very greatly
developed crests, a wider forehead with greater distance between the
tips of the supraorbital processes, and sharper constriction of the brain-
case behind the supraorbital processes.

Sexual differences in the skull are manifested only in somewhat
smaller average dimensions of female skulls. Age changes in their
skulls are manifested in the same way as in males.

The length of the intestine of adult Middle Russian wolves (2
specimens) is 460—575 cm, that of the juveniles 7-8 months old is 390—
420 cm (3 specimens); the ratio to body length in the former is 4.13
and 4.62, while in the latter, it is 3.64 and 3.86. The relative weight
of the heart (Hesse index) varies from 7.32 to 13.07; in young it is
apparently less than in old (Heptner and Morozova-Turova, 1951). The
length of the intestine (without the caecum) and the corresponding
index of two adult males from the extreme north (tundra of Arkhangel’sk
district—Taimyr) are 698 cm and | : 5.3 and 490 cm and | : 4.0. Their
heart weight is 800 gm and 16.4% (?) and 437 gm and 9.34 % (from
material of V. Makridin).

The diploid number of chromosomes is 78, the fundamental num-
ber, 80 (Matthey, 1949).

173

< “SA
НК RNA PRA, Ny

У ee

age

ety
HES

Fig. 47. Skull of wolf, Canis lupus L. Sketch by N.N. Kondakov.

129

174

The dimensions of wolves are subjected to geographic variation.
Body length of an adult wolf ranges between 105 and 160 cm; tail
length between 29 and 50 cm (usually from 40 to 50); hind foot length
is about 220—250 mm; and ear height is about 110—190 mm. Height at
the shoulders is 80-85 cm, and it may reach 100 cm.

Weight of adult Middle Russian wolves usually fluctuates between
32 and 50 kg. Females are not usually so massive as males, of some-
what smaller dimensions, and their weight is considerably less than
males. Average weight of females constitutes about 80-85% of average
male weight.

Information on the weight of wolves available in the literature,
especially old hunter’s and popular tales, are exaggerated. This is ex-
plained by the fact that they are mostly based on the determination of
the weight of particularly large animals “by eye”. Recently, in some
districts, mainly in the middle zone of the European part of the USSR,
accurate data on the weight of large series of animals appeared (see
section on “Geographic Variation”). These figures make us deal cau-
tiously with many of the old data on the average weight of wolves from
different parts of the range.

Nevertheless, huge wolves are sometimes found in some places.
Such animals, however, are very rare. In Middle Russia in general
form, the maximum weight of wolf is given as 69—79 kg (Ognev, 1931)
and 80 kg (Zvorykin, 1939).

Separate more accurate cases, recently verified, are the following:
In Saratov district, a wolf weighing 62.4 kg was recorded (Ognev,
1931); in the forest zone of the European part of the country, 69 kg
(A.A. Sokolov); in the districts around Moscow, a male weighing 76
kg is known—this was the largest one among 250 animals killed by the
famous wolf-hunter V.M. Khartuleri (Heptner and Morozova-Turova,
1951). In the Ukraine, animals of 92 kg (Lugansk district) and 96 kg
are known (Chernigov district; Korneev, 1950; circumstances under
which weight of these animals were determined are not known). In the
Altai, a male weighed 72 kg (Afanas’ev et al., 1953). In the Zoological
Museum of Moscow University, there is a mounted Middle Russian
wolf (from Е.К. Lorentz), which weighed about 80 kg (“5 poods’”)
(V.G. Heptner).

Information from Vladimir district in recent years indicates that
here there is a relatively high percentage of large animals. Of 641
wolves caught during the years 1951—1963 inclusive, 17 animals which
attracted attention because of their particularly large size were weighed.

132

175

They were caught in Petushinsk, Muromsk, Sobinsk and Suzdal’sk
regions and had the following weights (kg): Males—48 (February), 49
(January), 52 (February), 52 (November), 56 (February), 65 (Decem-
ber), 68 (January), 70 (March), 76.3 (March), 79 (January);
Females—40 (March), 41 (February), 45 (February), 48 (November),
55 (December), 58 (March), 62 (January). Along with these, three
adult wolves (older than two years), but from “the smallest” were
weighed: males 32 and 36 kg (March, April, Gorokhovetsk region) and
a female 30 kg (March; data of hunting inspection of the district—N.D.
Sysoev).

Concerning the Altai animal, it is known that it had “little meat”
in the stomach, but no other information was given. Sometimes the very
large weights of individual animals are related to the fact that they
were supposedly weighed with their stomach quite filled with food;
recently some have averred that a large wolf can eat 10—15 kg (!) of
meat. Just now this has been clarified, and information about the vo-
racity of wolves is отеайу exaggerated. Of 115 Voronezh wolves, only
one had 2 kg of meat in the stomach, while all the rest had much lesser
amounts (Merts, 1953). Of nearly 50 Saratov wolves, none had more
than 3 kg of food in the stomach (V.G. Heptner). Therefore, the weight
of some animals under all conditions very greatly exceeds the extremes
of normal variation (V.H.).

Systematic Position

Within the genus Canis, which is considered here in its most narrow
sense, C. /upus must be considered a more specialized progressive
form. Its morphological specialization as a predator is, apparently,
related to its adaptations for hunting large animals. Apparently, with
this, i.e. with its group way of life in the course of the greater part of
the year, are also related the progressive, highly developed, psychic
characters of the wolf (the structure of the pack, the capacity for
domestication, and so on). Obviously, we must evaluate the systematic
position of the wolf in the widest interpretation of the genus (see
characteristics of the genus and family). (V.H.).

Geographic Distribution

Almost all of Eurasia (except the extreme south and southeast) and
North America (except the extreme south).

176

131 Fig. 48. Sole of front foot of a wolf caught in the Sobinsk region of Vladimir
district. Winter fur. Natural size. Live weight of the animal 79 kg. Sketch from
nature by A.A. Alyapushkin.

134

177

The range in the USSR constitutes a very great part of the species
range and occupies nearly the entire territory of the country.

Geographic Range in the Soviet Union

The wolf is absent only on the Solovetsk islands, Franz-Josef Land,
Severnaya Zemlya and the Karagin, Commandor and Shantar islands.
Sakhalin is also included in the range (Shrenk, 1858; A.M. Nikol’skii,
1889, Suprunenko, 1890); however, since the beginning of present
century, it does not live there, and only occurs occasionally as a tran-
sient, crossing Nevel’sk Strait on ice. Wandering animals do not become
established (V.G. Voronov and A.G. Voronov).

Concerning the Kuril islands, information in the literature is in-
definite and contradictory and, to a considerable degree, erroneous.
Wolves were recorded on Kunashir and, with doubt, on Iturup
(Kuznetsov, 1949) and Paramushir (Podkovyrkin, 1950); wolves re-
corded on Shumishu (Sergeev, 1947) are simply feral dogs (Podkovyrkin,
1960). According to the most recent data, in the middle of the 60’s, no
wolves were found anywhere on the Kuril islands, and all records
pertain to feral dogs, which, in particular, are present on Iturup, Urup,
Shumishu, Kunashir and Paramushir (V.G. Уогопоу).

In the north, wolves are met with on Kolguev, Vaigach, and South
islands of Novaya Zemlya, on White Island, on the coastal islands of
the western (Nordenskiold island), northern and northeastern Taimyr
(in particular, Samuil island), on the Lyakhovsk islands and on the New
Siberian archipelago—on New Siberia and Kotel’n (E.I. Shereshevskii)
and Wrangel’ island.

At the extreme limits of its range in the north, in particular in the
northern Taimyr and on the islands to the east of Novaya Zemlya, the
wolf occurs irregularly and as a transient, not even every year. Thus,
on Wrangel’ island only one migration by three animals was known in
1938 (Manteifel’, 1947). Since migrations here of foxes are known,
this instance is probable although the width of Delong strait is more
than 150 km.

In the Arctic zone, the distribution and intrusions of wolves are
connected with the distribution and migrations of domestic and wild
reindeer (see reindeer in vol. I). This applies to individual years as well
as to considerable periods of time. Thus, wolves which fairly regularly
inhabited the southern island of Novaya Zemlya have disappeared with
the disappearance of the wild reindeer from this region and have not
been met with for a long time there (see “Biology” section).

‘iomdaH “D'A “USSN oy} ш “TY sndiyz siudD ом dy) JO UONNGMISIP oY) JO SoLepuNog илэ]еэ рие шэциом ‘6p “SIA 1

о , р 0 o8 a i
<. Yl >
5 i cig va
\ : J
me К р А = ry core

136

Wy)

To the south of the designated northern boundary, the wolf is
encountered everywhere, including the extremes of high mountains and
deserts, though in very different numbers. The wolf is extremely rare,
and absent in some places over considerable areas only in the taiga,
which owing to its heavy snows and the relative poverty of food re-
sources, represent the most unfavorable situation for the animal’s life.
When living conditions change, the wolf also occupies these districts.
Thus, in the north of Arkhangel’sk district, in connection with clearing
of forests, the spreading of stock-raising to the north, an increasing
number of forest wolves are nowadays energetically moving northward
into the taiga districts which were previously unavailable to them
(Parovshchikov, 1959). In the last decades, due to felling, road con-
struction, appearance of villages and similarly due to the disappearance
in many places of its enemy—the tiger—the wolf occupied a consid-
erable part of Ussuri taiga where it was previously absent and is
quickly spreading northward (Abramov, 1954 and others).

A long time ago wolves were completely destroyed in the Crimea
and were totally absent there for a long time. They appeared there
during the Civil War, but they were again eliminated. During the Sec-
ond World War and the first years thereafter the animals appeared
again on the peninsula, but quickly disappeared.

Wolves have migrated across the ice to several islands in the
northern part of the Caspian Sea, and at the time of seal hunting,
attracted by the abundance of food (carcasses of killed animals), they
go far onto the ice of the open sea (from the eastern shore, V.G.
Heptner).

Geographic Range outside the Soviet Union

This comprises Japan (now extinct), the Korean peninsula, China south-
ward to the Yangtze-kiang, the Mongolian Republic, Tibet, the Hindustan
[Indian] peninsula to Bengal on the east and at least to Dharwar (about
latitude 15°30' N. Lat., a little east of Goa) to the south, Baluchistan,
Afghanistan, Iran, Iraq, the Arabian peninsula, the countries of the
Levant and all of Europe. In Europe the wolf is exterminated in Ireland,
England, France, Holland, Belgium, Denmark, Switzerland and in all
of Central Europe. In the German Democratic Republic and even in the
Federal Republic of Germany during the war and the post-war years,
the appearance of few vagrant individuals was noted.

134

180

Fig. 50. The boundaries of distribution of forest wolf (solid line) and tundra wolf
[broken line]* in Arkhangel’sk district. Arrows indicate direction of winter
movements of tundra wolf to the south (after Parovshchikov, 1959).

In North America, the range of the wolf (reconstructed) occupies
nearly the entire continent; on the south, to the southern border of the
Mexican highlands (near 20° N. Lat.), and on the north, the whole
Arctic archipelago. It also occupies the northern coast of Greenland and
the western and eastern to 69° N. Lat. (Disko [Island], Godhavn and
Scoresby Sound). Accordingly, the most northern point of the animal’s
occurrence is located only 700 km from the pole.

In the south, the range does not include the extreme margins of the
western coastal parts of the continent, beginning at 41° N. Lat. in the
south, both in the United States and in Mexico, and the coastal areas
of eastern Mexico to the Rio Grande. The range also does not include
the Queen Charlotte islands (it is on Vancouver Island) and the islands
of the Bering Sea (although they are known to have crossed the ice to
Saint Lawrence Island). The wolf is exterminated in a considerable part
of the United States and several races are completely extinct. (V.H.).

*Not in original Russian—Sci. Ed.

(Y¥-

Aw }

о > . м
ts

ae:
1 о.
ae
Ae [9--
ov ЧА

S { р,
S (7

fil pire <<

182
Geographic Variation

The huge range and their occurrence in quite different natural condi-
tions permit us to infer that wolves show considerable geographic
variation. Authors of the past century have pointed out the formation
of races in wolves. This is confirmed in our own time by many data.
However, geographic variation of the wolf is, so far, not satisfactorily
studied. This applies to the Old World as well as the New where, in
particular, many improbable subspecies were distinguished. However,
analyses of several of our populations were made at a good level in
recent years. These analyses are, however, very few and concerned
only with craniology. Individual variation in color is quite great, and
wolf skins are scarce in museums because of their high price. There-
fore, one may attempt to characterize at the present moment only a few,
better distinguished races of our fauna, and refer only to what may be
outlined. This review of the forms and characters given below thus
bears a preliminary character.

1. Tundra wolf, С. [. albus Kerr, 1798 (Syn. turuchanensis,
kamtschaticus, dybowskii).

A wolf of large size. Body length of males (23) 118—137 cm, tail
length 42—52; average weight 40 kg, maximum 49 (1 specimen of 55,
with the stomach contents). Body length of female wolves (23) is 112—
136 cm, tail length 41-49 cm, average weight 36.6 kg, maximum 41 kg.

Greatest length of skull of males (10) 248.5-+(M 257.2)-270 mm;
females 237.5—М 247.9 mm)-256.4 mm.

Pelage is very long, dense, fluffy and soft. Length of top hairs is
150—160 mm, guard hairs, 80—150 and underfur, about 70. The usual
color is very light and gray; underfur has two zones of color: the lower
is lead-gray, the upper is reddish-gray.

Concerning the general light color tone, two main types are found, |
connected by intermediate forms: light-gray with reddish shade and
bluish-gray without this shade. The latter is characteristic of juvenile
wolves; old animals are lighter in color and have a reddish frosting.
Moreover, the color of the skin changes noticeably during the course
of the winter. In November—December, wolves as a group are darker
than in April-May, when the fur strongly fades under the influence of
the spring sun. Very light individual wolves are sometimes found in
May.

Dimensions and color description apply to wolves of Malozemel’sk
and Bol’shezemel’sk tundras (Nenets Nats. Okrug, Arkhangel’sk dis-
trict; Makridin, 1959).

183

Dimensions of adult Taimyr wolves (caught in 1961) are as fol-
lows. Body length of males (11), 123-(M 127.7)}-146 cm; of females,
112—(М121.3)-129 cm; tail length of males, 42-(M 44.4)48 cm, of
females, 40-(M 42.6)—45 cm; hind foot length of males, 25—М 26.0)
29 cm, of females, 23-(M 25)-27 cm; ear height of males, 11~M
12.7}-13 cm, of females, 114M 12)-13 cm.

Greatest length of skull of males (6), 258.0—(М 267.0)288.3 mm,
of females (7), 239.44M 251.6)-261.4 mm; condylobasal length of
skull of males, 238.0—(М 247.6 mm)-—*, of females, 221.6—(М 232.3)-
243.0 mm; zygomatic breadth of males, 134.3—М 146.9)-160.7 mm,
of females, 132.5—М 137.1)-142.2 mm; breadth above canines of
males, 47.0—(М 51.1)-55.4 mm, of females, 41.4-(M 45.7)-47.3 mm.

Data on maximum weight of northern wolves were obtained from
nearly 500 animals caught during 1951—1961, on the Taimyr and west-
ward to Kanin. The greatest weight was of an old male killed on the
Taimyr at the mouth of the Dudypta river—52 kg. Two animals, killed
on the tundra east of Kanin and on the Yamal, had weights of 48.8 and
46.7 kg (without stomach contents) (У. Makridin).

Found in the tundra zone and the forest tundra of the European and
Asian parts of the USSR and in Kamchatka.

Outside the boundaries of the USSR, perhaps in the extreme north
of the Scandinavian peninsula.

The belief that the wolves of the extreme north (tundra wolves) are
the largest race of Old World and are larger than the forest wolves is
quite widespread. Actually, tundra wolves are, apparently somewhat
smaller than forest wolves (Makridin, 1959 and others). Statements
that in tundra wolves “weight reaches 70 kg and sometimes more”
(Kuznetsov, 1952) as a characteristic feature of the population and the
subspecies is exaggerated. However, among tundra wolves, specimens
of more than 50 kg in weight are found. Moreover, it is possible that
the figures given are somewhat reduced, as they were obtained in a
locality where intensive hunting of wolves was carried out over the
course of several years by aircraft (see later).

As a whole, the tundra wolf is a well-characterized form, and
noticeably differs from the forms distributed in the forest zone by its
color and, apparently, its dimensions. At present it is impossible to
judge whether the wolves in all the area of its distribution are identical
or not. Concerning color, wolves of the northern Kola Peninsula are on

*Translator’s remark: a figure is missing in original Russian.

138

184

the whole somewhat different, and are more similar to forest wolves
and wolves of extreme northeastern Siberia. It is not excluded that
there are also some differences in the average measurements of the
various tundra populations. They have not been studied, but they do
not, apparently, exceed the limits of local populations within the bound- |
aries of the subspecies. In the Old World there are no populations of
almost pure white wolves similar to those living in the extreme north
of America (С. [. bernardi, С. |. manningi and others of the tundrarum
group).

This subspecies coincides with the polar assemblage of fur-bearing
animals (see beyond).

2. Middle Russian forest wolf, С. /. Jupus Linnaeus, 1758 (syn.
communis).

A wolf of very large dimensions, probably maximum within the
boundaries of our country.

Color of wolves of this race corresponds to that given above in
“Description” (p. 166).

Found in the forest zone of the European part of the country,
northward to the northern limit of forest, and southward including the
forest steppe. The exact limits of distribution to the south are not
known at all; to the west it is to the state boundary and the distribution
in the east is not established. It probably spreads more or less far into
Siberia, and possibly, occupies western Siberia.

Outside the USSR—in Europe except, evidently, the extreme
south.

The Middle Russian forest wolf is well differentiated from the
tundra wolf, although in the north, both forms penetrate into the area
of occurrence of each other (see “Geographic Distribution”).

It is usually considered that the weight of adult males is 40-45 kg,
a yearling, about 35 kg and juveniles about 25 kg. Maximum weight of
the Middle Russian wolf is considered to be from 69 to 80 kg (see above).

Dimensions of animals from separate regions are as follows: weight
of adult wolves of northern part of Ryazansk district (south of
Meshchera—a little south of the latitude of the Pra river) varies from
32 to 55 kg (based on 173 individuals); weight of the largest male was
56.3 kg (Kozlov, 1955).

Of 65 wolves killed in the region of Mordovsk preserve (Temnikov),
only one old male with worn teeth weighed 49 kg. The remaining adult
males had weights from 40 to 45 kg; adult females, from 30 to 35 and
only one female weighed 38 kg (Kozhevnikov, 1953).

139

185

The wolves of Voronezh district, 1.е. the forest steppe zone, appar-
ently are somewhat different from Moscow wolves of the true forest
zone and have the following measurements and weight. Average body
length, based on 154 animals killed in the region of Voronezh preserve,
was 125 cm in males and 122 cm in females. The majority of animals,
both old as well as yearlings, had a weight from 30 to 40 kg. Many
juvenile wolves reached a weight of 23 kg by the end of November.
Weight of adult (full-grown) males (48) averaged 40.0 kg; weight of
adult (full-grown) females (31) was 3142.1 kg, averaging 36.0 kg.
Weight of yearling male wolves (up to two years old) (8) was 26.6—
34.0 kg, averaging 32.0; weight of yearling female wolves (24) was
23.0—32.8 kg, averaging 29.0 kg (weight without stomach contents;
Merts, 1953).

Body weight of juvenile and yearling males in Belovezhsk Forest
varies from 23 to 35 kg; of females—from 22 to 29 kg (Table 9*).

The described race is related to the largest ones and, probably it,
and not the tundra wolf С. [. albus, is the largest form of Old World.
Beyond that, the figures given show that the average measurements of
the Middle Russian “forest wolf’, usually obtained from especially
large animals, are often exaggerated. Thus, out of more than 500 ac-
curately weighed animals, only 3 males had a weight exceeding 50 kg,
one of these from the Oksk preserve at 56.5 kg (V.V. Kozlov).

Within the defined region, wolves are not entirely of one type.
Thus, animals from the extreme west (Belovezhsk Forest) are, as shown,
somewhat smaller than from Moscow and, in general, those of the
districts around Moscow, but they do not differ in color. Wolves of

Table 9. Dimensions and weight (without stomach content) of adult wolves of
Belovezhsk Forest (Gavrin and Donaurov, 1954)!”

min. max. average min. max. average
Body length 105.0 140.0 119.0 97.0 124.0 111.0
Tail length 33.0 50.0 41.0 29.0 45.0 40.0
Hind foot length 20.0 28.0 25.0 21.0 26.0 23.0
Ear height 11.0 19.0 13.0 11.0 12.0 13.0

Weight 23.3 44.8 34.8 21.0 36.0 29.2

*In Russian original, 10, ап error—Sci. Ed.

"The authors did not give the number of males and females of their series. The
figures given here are based on the authors’ indication of sex ratio in the population they
studied.

186

61€l O7%PI 0601 SE 6351 05591 6611 Or yypesiq эвешозА7
€ tp OLb 9°8€ 9€ ГА L'9b ULE 6£ sasseooid jeyqioeidns pulyoq yypeolg
719 O'7L 0'55 98 6°79 vol 0'8r Or sassacoid yeyquovidns ye чуреэла
€ bP 8'05 Ce 6£ 6S aS p'8e СР [NYS JO Ч3реэ44 [21920123]
Lvl Z'06 5°69 Or "58 0°56 07 СР souog |езеи иээм1э4 onIns Jo WSUS]
0°66 СИГ 39°98 6$ 601 0 $96 $ aulued YIM м0443003 Joddn jo ц38иэ7
осо» бис *0'L0C Ze IFLET & 0662 0315 Se [INYs JO 438иэ| [езедо1Арио)
Оса. 0692. “0°17 Se 6966 — 05/0 = 01655 Iv [INys JO ysua] 15936910)
эЗелэле ‘хеш ии и эЗелэле = “XBW "и и

Эш ЭГ 81

35940 YSYZAAOJIG JO зэл|ом зпре JO ззиэшолизвош [INAS “Ol э19Ет,

8el

187

Voronezh district are, apparently, not smaller than those of the more
northern districts, although they are somewhat grayer and lighter in
color than those of Moscow, the color of which is more reddish. Among
145 winter skins (mainly from Voronezh preserve) 79% were gray,
10% light-gray and only about 20% consisted of skins that had “rusty
and bright-rusty” tone (Merts, 1953). It is probable that the lighter
color is generally characteristic of wolves of the forest-steppe region.
This may be the so-called “Central steppe wolf” of furriers (see beyond
p. 195). The Mordovsk wolves correspond to the Moscow ones in color
and measurements. Difference in measurements in the relatively small
areas mentioned may not represent geographic variation: in places where
hunting is very intensive, wolves have little possibility of living to full
development and attaining maximum size, and the proportion of the
relatively young animals in the population is large.

No comparison was made between our wolves and Scandinavian.
However, judging by the characters of animals from our northwest,
their assignment to the nominal form is correct.

3. Siberian forest wolf, С. /. altaicus Noak, 1911.

A large wolf, not less in size than the Middle Russian, С. /. Jupus.

Coloration is light gray, darkened along the middle of the back by
dark guard hairs. Yellow (ocherous) tones in skin color are absent or
weakly developed. The fur is dense, long and soft, but it is slightly
shorter and not so silky as that of the tundra wolf. Length of guard hairs
on the withers is 9—11 cm.

Found in Siberia and the Far East except the tundra zone,
Kamchatka, Trans-Baikaliya and Ussuri territory, and also, probably,
the western parts of Siberia; northern Kazakhstan.

Outside the USSR, found in the northern montane taiga of the
Mongolian People’s Republic.

The systematics of Siberian wolves is not worked out at all, al-
though wolves in the great territory extending from Urals to the Pacific
Ocean and from the tundra to the southern mountain ranges are hardly
identical. According to one view, the area eastward to Lake Baikal is
inhabited by the nominal form distributed in the European part of the
Soviet Union (Flerov, 1935), and according to another (Kuznetsov,
1952), it is the “Siberian forest wolf’ which is a form different from
the Middle Russian and occupying all of Siberia except its tundra parts,
and northern Kazakhstan.

Until clarification of geographic variation in Siberian wolves, this
form is conditionally separated. The name given here is also condi-

140

188

tional, it being, apparently, the only one applicable to the Siberian forest
wolf. Since the western Siberian and northern Kazakhstan wolves are,
apparently, very close to the Middle Russian ones, then the name С. /.
altaicus is, above all, applied to the Middle- and East Siberian animals.

4. Steppe wolf, С. /. campestris Dwigubski, 1804 (syn.
arundinaceus).

Average dimensions are somewhat less than the Middle Russian
wolf, С. /. lupus. Pelage is shorter, coarser and sparser. Color of the
sides is relatively light, gray, on the back rusty-gray or brownish with
a quite strong admixture of black hairs.

Found in the Pri-Caspian steppes, steppes of the Cis-Caucasus and
the lower Volga, the steppes of Kazakhstan (except the north?), and the
steppe region of the European part of the Union.

Outside the USSR, perhaps in the steppe section of Romania and
Hungary (in the past).

Apparently, this is the wolf of the Kazakhstan steppes (except the
north), intermediate between the northern forest and southern desert
wolves, which also penetrates into the steppes of the Cis-Caucasus and
the European part of the Soviet Union. However, it is a form of weakly
determined characters and distribution. The differences between steppe
wolves and Middle Russian ones have been mentioned and are accepted
by all authors, and they obviously exist, although there has been no
study of this question based upon reliable material. At the present time,
the wolf is absent or very rare in considerable areas of our European
steppes. In the northern Ukraine and to the west, also in the steppe
lives, apparently, not the true, relatively small, steppe wolf, but the
larger forest or “forest-steppe” form. It is probable that the latter colo-
nized the western parts (Odessa, Khersonsk, Nikolaevsk districts), where
the true steppe wolf was almost completely exterminated during war
and in the post-war years (Korneev, 1950).

In the population of steppe wolves, particularly in the Ukraine, few
very large individuals are encountered (see above). They are mainly
found in the northern part of the steppe zone. Apparently, these indi-
viduals intrude from the north (see beyond).

С. 1. campestris, apparently, corresponds to the “Caucasian steppe
wolf” (see later).!°

'3In recent years, many references deal with the problem concerning the exist-
ence in the past in the steppes of Hungary of the very small “reed wolf’, sharply
distinguishable from the large Carpathian forest wolf. According to the view of some

189

Note: With regard to the type locality selected here (Cis-Caucasian
steppes) (see synonymy), the following must be taken into consider-
ation. Dwigubsky states “in desertis ad mare Nigrum et Caspium
interjacentibus, Kirgisicis, Bucharicis, ad f. Yeniseam, ubi, dantur colore
sordide flavorubentes” (1804), i.e. “In desert places situated between
the Black and Caspian seas, in Kirgiz steppes, in Bukhara, and at the
Yenisei river where dirty-yellowish-reddish forms are encountered”, or
“in the steppes near the Caspian and Black seas; in Kirgiz steppes and
near the Yenisei river, where red wolves also occur” (1830).

In all the districts outlined by Dwigubsky, wolves are undoubtedly
not identical. In accordance with the beginning of the first text given,
and in connection with the unclear systematic position of wolves in the
west of our steppes (Ukraine), it is here suggested to restrict the very
wide definition of Dwigubsky to the mentioned terra typica restricta.
Dwigubsky [Dvigubskii] moreover stated (1830) that “in my traveling
in the steppe between the Dnepr and the Don, in 1805 (i.e. after the
publication of the description. V.H.), I had occasion to see such wolves
even in the Bakhmutsk district near the Donets.”

The suggestion about the affiliation of the name campestris to the
jackal (Pocock, 1941) has no scientific basis; in almost all the territory
mentioned by Dwigubsky, the jackal is absent and was never previ-
ously found.

5. Caucasian wolf, С. [. cubanensis Ognev, 1922 (syn. hajastanicus).

Wolf of medium dimensions (less than Middle Russian). Winter
coat is poorly furred, coarse, short and quite sparse. Length of guard
hairs on the withers does not usually exceed 70—75 mm. Coloration is
dirty, dull gray-ocherous, quite dark due to the more or less evenly
distributed guard hairs on the skin. Greatest length of skull of males
240—272 mm, of females 224—251 mm; condylobasal length of skull of
males is 221—250 mm, of females 209—234 mm; zygomatic breadth of
skull of males 128—152 mm, of females 11*—132 mm.

Found in the Great Caucasus and Trans-Caucasus.

Outside the USSR it is not known; probably spreads into the con-
tiguous territories of Iran and Turkey.

authors, which is, apparently, justified, the “reed wolf’ is simply the jackal. It is not
excluded that our steppe wolf described above might have penetrated quite deeply to
the west. It is not, however, similar to that described by some authors (Nad) under
the name “reed wolf’ (Rohrwolf).

*Error in original Russian text—Sci. Ed.

141

190

The distinctiveness of Caucasian wolves is mentioned by a series
of authors and is, evidently, real. However, the form is very poorly
studied. Its distribution is also little known. Moreover, in various parts
of Caucasus, wolves are, apparently, not identical. Thus, on the plains
of the Trans-Caucasus, the animals are somewhat smaller than those
inhabiting the northern Caucasus. It is possible that small animals
belonging to the group of desert wolves (desertorum-—pallipes), de-
scribed from Asia Minor, penetrate here from the south. Wolves of the
Armenian highlands, described as С. /. hajastanicus, are somewhat
larger than those of the plains (they correspond, in dimensions, to those
inhabiting the Kuban Caucasus) and have softer, longer fur (Dal’,
1951; Vereshchagin, 1959). However, their separation as an indepen-
dent form is insufficiently argued.

6. Desert wolf, С. 1. desertorum Bogdanov, 1882 (syn.
bactrianus).

Measurements are considerably less than those of northern forest
wolves; the smallest form of those inhabiting our country.

Coloration is light, gray-sandy or yellowish-gray with a few black
guard hairs on the upper part of the body, mainly along the spine. In
the area of the occiput and the outer side of the ears, reddish tones are
developed. Winter fur is not very luxuriant; quite sparse and coarse.
The tail is poorly furred.

Weight reaches 35—40 kg.

Found in plains areas, including the deserts of Middle Asia and
southern Kazakhstan north to the middle Emba, and the northern Pri-
Urals and in the northeast to the region of the Chuya river, Betpak-Dala
and Pribalkhash’e.

Outside the USSR, found in Afghanistan and Iran, at least in the
northern parts of these countries.

The desert wolf represents a well-differentiated form. Specific,
precise data on the dimensions of this form have not been published,
although its small measurements are indicated by all authors. At the
same time, large individual specimens are encountered in the wolf
population of Turkmeniya (Heptner, 1956).

The wolf of the Trans-Caucasian plains and the Mongolian Repub-
lic has much in common with the desert one. С. /. desertorum 1$,
evidently, closely related to the Indian wolf, С. /. pallipes, although this
form is, apparently, still smaller and also somewhat differentiated by
coloration and in fur characteristics. The boundary between the geo-
graphic ranges of the forms desertorum and pallipes is unknown.

142

191

This form, apparently, corresponds to “Middle Asian steppe wolf”
of the Middle Asian bloc of furriers (see beyond).

7. Tibetan wolf, С. [. chanko Gray, 1863 (syn. [ашеег auct., nec
laniger Smith).

Dimensions comparatively small, noticeably smaller than С. /. Jupus,
but slightly larger than С. /. desertorum.

The coloration is light, whitish-gray with an admixture of darker
brownish tones on the upper part of the body. Along the spine there is
a well-defined dark stripe. The winter fur is soft, long, quite luxuriant,
with a well-developed undercoat. The tail is densely furred with long
hairs.

Weight, apparently, does not exceed 45 kg.

Found in the Pamirs proper and the Pamir-Alai system and also the
Tien Shan.

Outside the USSR, it is found in montane regions of Central Asia,
Dzhungariya and Tien Shan, across Tibet to Gansu, Sichuan and Yunnan
in the east and Inner Mongolia in the north.

This well-defined race corresponds to the “forest (mountain) wolf”
of the Middle Asian fur trade.

Note: As regards the name and type locality of this form (see
synonomy) the following must be taken into consideration. “Chinese
Tatary,” from which it was described, is extremely ill-defined. How-
ever, none of the authors concerned with the nomenclature of wolves
(Allen, 1938; Pocock, 1941) attempted to specify it. This form has been
given a very wide distribution in Central Asia “from Chinese Turkestan
and the Tien Shan through Tibet into Mongolia, northern China, Shanxi,
Sichuan, and apparently, Yunnan; the western Himalayas in Kashmir,
at least from Chitral to Lahul” (Pocock, 1941). The uniformity of
wolves in these regions and above all the identity of the animal of the
low desert of Kashgariya (Chinese Turkestan) and high montane Tibet
and the Himalayas is very doubtful. Since the long furred wolf is, first
of all, associated with Tibet, and since the name chanco is employed
in place of the name /aniger to Tibet animals, it is here suggested that
the terra typica of the form chanco be considered Tibet.

Within the borders of the Kazakh republic, therefore, the ranges of
four races of wolf come together. In the north (still 100—130 km south
of Kustanai, V.G. Heptner) and in the extreme east, the large “Siberian
[and Middle Russian] forest wolves” (С. [. lupus and С. /. altaicus)
live. In the south is distributed the small, sandy-colored desert wolf, C.
l. desertorum. The entire intervening region is, apparently, occupied by

192

a form transitional (intermediate) between them. It evidently follows
that it should be considered C. /. campestris, the range of which passed
through the region north of the Caspian Sea westward to the southern
Russian steppes. In the steppes of Kazakhstan, far beyond the region
of its usual occurrence, isolated individuals of both the southern and,
especially the northern forms, wander. Wolves of the northern type
have been noted at middle Emba (Afanas’ev et al., 1953). In Tien Shan
the montane form, C. /. chanco lives. There is insufficient basis to
assume the existence in Kazakhstan of two undescribed subspecies
(Afanas’ev et al., 1953).

8. Mongolian wolf, С. [. tschiliensis Matschie, 1967 (syn.
argunensis, dorogostaiskii).

Dimensions are not large—like С. |. desertorum, or somewhat
larger, but markedly smaller than those of the Siberian forest wolves.

Coloration is dirty gray, frosted with a weak admixture of ocherous
color and without bright pale-yellow and chestnut tones. The fur is
coarse and stiff.'*

Total body length of males (18), 93-(M 115)-158 cm; tail length,
30—40 cm; hind foot length, 16—24 cm; ear height, 10—14.5 cm; shoul-
der height, 58-89 cm. Weight, 26-(M 31.3)-37 kg. Total length of
females (8), 90—(М 99.6)-109 cm; tail length, 30-40 cm; hind foot
length, 16—23 cm; ear height 9.5—13 cm; shoulder height, 57—75 cm.
Weight, 22—(М 27.7)-30 kg (animals from southern part of Ussuri
territory; G.F. Bromlei).

Greatest length of skull of males (5), 226.0—(М 2393.7)-253.5;
condylobasal length of skull (3), 224.0—(М 229.8)—236.5; interorbital
breadth (5), 38.5—М 42.0)-45.5; postorbital breadth (5), 38.5—М
40.5)—43.5; zygomatic breadth, (5) 114.0—(М 126.7)-139.5 (speci-
mens from eastern Trans-Baikaliya; Skalon, 1936).

Found in eastern, southern and southwestern Trans-Baikaliya and
in the Ussuri territory.

Outside the USSR, in the Mongolian Republic, except the northern
montane and taiga parts, in Inner Mongolia and probably, on the Korean
peninsula.

The described form is, apparently, quite well characterized—tt is
the wolf of the eastern (Central Asian) section of the steppe and desert

'4Statements that the “eastern (Trans-Baikal) wolves differ by their long tails, in the
majority of cases curved upwards at the end” (Manteifel’ and Larin, 1949) cannot be
considered seriously.

143

193

zone of Eurasia. Based on its coloration and dimensions, it possesses
features relating it to С. /. desertorum. Generally, this form is still very
poorly studied. (Characteristics and distribution based on data of Skalon,
1936; Loukashkin and Jernakov, 1934; and G.F. Bromlei.)

Within the USSR, apparently only a narrow strip of the northern
edge of its range is found; the Siberian forest wolf lives around Lake
Baikal and, probably, around Chita.

The presence of this “steppe” form in the forests of Ussuri territory
is related to wolves from the south having reached there only in the
present century in connection with the clearing of taiga and the de-
crease in the number of their antagonist—the tiger. Formerly, wolves
were generally absent.

The nomenclature of this form is unclear. The name employed,
given all the inadequacies of description is, apparently, concerned with
it. At present there is no basis for suggesting that the wolves of Trans-
Baikaliya and northeastern China differ from those from Liaodun*.
Nevertheless, the name ¢shiliensis is considered conditional pending а
solid systematic revision of the wolves of Inner Asia and the Far East.

9.2? Sakhalin wolf, С. 1. hattai Kishida, 1931 (syn. rex).

The characteristics of this proposed form are unclear and its indepen-
dence cannot be considered established. Its description is unsatisfactory.
It was described from Hokkaido, where it is now exterminated. It was
applied to Sakhalin and “perhaps” to the Kuril islands (Harper, 1945;
Ellerman and Morrison-Scott, 1951). As shown above (see “Geographic
Distribution”), the wolf is absent on the Kuril chain, and on Sakhalin, it
has not been met with since the beginning of the present century. There-
fore, it is impossible now to decide which wolf lived on Sakhalin. Intrusions
of the Siberian forest wolf occur from the mainland across Nevel’sk
Strait on the ice but, apparently, it does not remain.

* * *

Geographic variation in the wolf finds its reflection in the sorting of
skins into groups, as is usual in the fur trade and this attracted the
attention of zoologists. The value of these data lies in the fact that they
are based on the examination of large quantities—tens of thousands of
skins. In some cases, there is a quite exact coincidence of the groups
with subspecies, while in others, differences in classification are sub-
stantial.

*Place in northeastern Xinjiang province—Sci. Ed.

144

194

“Characteristics of winter-killed skins of different groups of wolves
are as follows:

Polar group: The entire tundra and forest-tundra belt of the USSR
and Kamchatka.

Skin size is very great. In spite of the thin flesh side of the skin,
large size of skins gives them a considerable weight—one hundred
skins weigh on average 150 kg. Weight of adult animal reaches 70 kg,
and sometimes more. Pelage is especially luxuriant, thick, and in par-
ticular, very soft. Length of guard hairs on the nape of winter skin is
11—13 cm. The abdomen is covered with long dense fur. The flesh side
of the hide is light and thin. Coloration of the fur is light or dark gray,
tinged with bluish. Wolves of the Siberian tundra have usually the
spine and the sides of light, pale-grayish color tinged with blue. The
dark guard hairs grow mainly on the spine, forming a longitudinal belt;
on the sides, dark tips of the guard hairs are not well defined. The
abdomen and lower thorax are whitish. The limbs are almost white.

In the extreme north of the Far East, wolves have similar color but
usually mixed with a yellowish tinge. Murmansk wolves have a darker
color due to the great development and distribution of black tips of the
guard hairs on the skin.

Siberian group: All of Siberia and the Far East except the tundra
regions, and also the northern part of Kazakhstan.

The group is subdivided into skins of the Siberian forest and steppe
wolf.'°

Siberian Forest wolf. Skins have the same measurements as those
of the polar wolf. Weight of one hundred hides is 148 kg. Weight of
an adult Siberian wolf reaches 70 kg and even more. Pelage is dense,
long and soft, but is of lower quality and not so silky as that of the
polar wolf. Length of guard hairs on the nape ranges from 9 to 11 cm.
Flesh side of the skin is compact and quite thick. Coloration of winter
fur is usually light gray, darkened along the spine due to dark tips of
the guard hairs. Yellow tints are either absent or poorly developed.
Abdomen is whitish. Limbs are gray.

'SDivision of the group into subgroups of forests and steppe generally reflects
the basic types of individual variation in coloration and fur quality. In some cases it
indicates the migrations (sometimes seasonal) of a certain number of animals of one
race into a region inhabited by another race. Sometimes, as mentioned above, these
groups correspond to particular subspecies.

Data on animal weight, as mentioned above, in the reviewed groups is some-
times overstated.

195

Siberian Steppe wolf. Skin size is as in the forest wolf. Pelage is
moderately luxuriant, but coarse. Coloration of the spine and the sides
is gray with clear brownish or reddish tints and with black tips of the
guard hairs on the spine well developed. Abdomen is reddish or yellow-
ish. Limbs are brown.

Kazakhstan group: Kazakhstan, except its northern regions.

Skins of the Kazakhstan group are also divided into skins of the
forest and steppe wolf, which are similar to the hides of steppe and
forest varieties of the Siberian wolf, but differ in their smaller dimen-
sions and less luxuriant pelage (weight of 100 hides of Kazakhstan
wolf is 112 kg, and length of guard hairs on the withers of forest wolf
reaches 7 cm).

Middle Asian group: Middle Asia.

Skins of this group are divided into skins of forest (more correctly,
montane) and steppe Middle Asian wolves.

Forest (montane) Middle Asian wolf. Dimensions are small, and
weight of 100 hides is not more than 100 kg. Weight of an adult animal
does not exceed 45 kg. Pelage is quite soft and long, but not particu-
larly dense. Coloration is light-whitish-gray, slightly darkened on the
spine by dark guard hairs. Side and abdomen are whitish.

Steppe Middle Asian wolf. Dimensions are still less than montane
wolves of Middle Asia. Live weight is usually less than 40 kg. Pelage
is not very luxuriant, coarse, coloration of spine is light yellowish-gray.
Abdomen is grayish.

Central (Russian) group: European part of the USSR, except tun-
dra regions, Ukraine, Crimea, northern Caucasus, and the lower Volga
area. They are divided into the forest and steppe central wolves.

Central forest wolf. Dimensions are large—nearly the same as
those of the Siberian forest wolf. Weight of 100 hides is 166 kg.
Weight of live animal is up to 70 kg. Pelage is of average luxuriance
and softness. Length of guard hairs on the withers is usually not more
than 9 cm. Coloration of fur in the majority of skins is quite dark, gray,
usually with brownish tints. The quite dark color of the skin is due to
there being black guard hairs distributed on the entire middle dorsal
part of the skin, somewhat darker in the region of the vertebrae. Ab-
domen is whitish or yellowish.

Central steppe wolf: Dimensions are slightly less than the forest
form. Pelage is not very luxuriant, coarse and bristly. Coloration of the
spine and sides is brownish or reddish-gray. Abdomen is yellowish.

145

196

Caucasian group: Ukraine, Crimea, northern Caucasus, Trans-
Caucasus, lower Volga area.

Skins of this group are subdivided into two categories: forest and
steppe Caucasian wolves.

Caucasian forest wolf (true montane. V.H.). Dimensions moderate.
Weight of 100 skins is 146 kg. Weight of live animal reaches 50 kg.
Pelage is not very luxuriant; sparse, coarse. Length of guard hairs on
the nape is not more than 7.5 cm. Flesh side of the skin is porous and
thick. Coloration of the spine and the sides is dirty-gray, usually quite
dark due to numerous evenly distributed dark guard hairs. Abdomen is
dirty whitish with yellow tints.

Caucasian steppe wolf. Dimensions are the same as those of the

forest wolf. Pelage is short, sparse, and coarse. Fleshy side of the skin

is thick, porous. Coloration of the spine is usually rusty-brownish or
rusty-gray with dirty tints, and is strongly darkened from dark tips of
the guard hairs. Sides are lighter in color. Abdomen is dirty-whitish.

Skins of the polar wolf are considered the most valuable—among
the remaining, the best are skins of the Siberian wolf; skins of the
Central and Kazakhstan wolves are less valued, and those of Middle
Asian wolves still less so; the poorest skins are those of Caucasian
wolves” (Kuznetsov, 1952).

From parts of the range lying outside the boundaries of our country in
the Old World, the following forms are recorded.

1) С. 1 signatus Cabrera, 1907—Тбепап Peninsula; 2) С. [. deitanus
Cabrera, 1907—Spain; 3) С. 1. italicus Altob., 1921—taly and Sicily;
4) С. 1. kurjak Bolkay, 1925—Yugoslavia; 5) С. 1. hodophylax Tamm..,
1839—Honshu, Hondo*, exterminated; 6) С. /. pallipes Sykes, 183 1—
India, Baluchistan, southern Iran, Iraq, ? Asia Minor, northern part of
the Arabian peninsula; 7) С. 1. arabs Pocock, 1934—-southern part of
the Arabian peninsula.

The existence of some forms in Europe (two in the Iberian Penin-
sula) is very doubtful, the remainder are, apparently, real (especially
С. 1. pallipes).

*Honshu and Hondo are synonyms, the first being the modern, the second the old,
name of the main Japanese island—Sci. Ed.

146

197,

Geographic variation of wolves in the Old World, as regards char-
acteristics of size and also color is quite regular. The largest wolves are
northern, in the tundra and forest zones.

Steppe wolves are somewhat smaller in size, as in wolves of the
Caucasus, and wolves of the deserts of Middle and Central Asia are
still smaller. Wolves of this type also inhabit the mountains of Central
Asia; if some of them are larger than those of desert plains, the differ-
ence on the average is usually insignificant. The smallest wolves inhabit
the southern margins of the range—India, Baluchistan, Iraq and the
Arabian peninsula'®. Wolves of types desertorum, tschiliensis and
chanco comprise a transitional group from the large northern to
the small southern ones. Accordingly, Bergman’s rule is quite clearly
demonstrated.

Wolves with the lightest color are those of the extreme north—the
tundra zone; more southerly ones living in the forest zone are more
intensely colored, while those of the extreme south—the desert zone,
have light ocherous “desert” color. Animals of the mountains of Central
Asia are somewhat more intensely colored than in the desert. Accord-
ingly, Gloger’s law is also demonstrated.

Geographic variation in New World wolves is well demonstrated.
Twenty-six well-founded forms are usually accepted (Young and
Goldman, 1944; Miller and Kellog, 1955; Haltenorth and Trense, 1956;
Hall and Kelson, 1959)!’. Both a priori and from the published mate-
rials, it is sufficiently clear that this number is overestimated and the
number of actually existing and the really differentiated subspecies of
American wolves is considerably less. Nevertheless, there is no doubt
that geographic variation of animals on this continent is clearly ex-
pressed and significant. There is a quite definite geographical trend
related both to size and color.

The largest wolves live in the North; towards the south they gradu-
ally become smaller, having minimal dimensions in the south of the
range. The largest wolves are in the extreme northwest—in Alaska and
the Mackenize basin (pambasileus, mackenzii, tundrarum). Here ani-
mals attain a weight of almost 80 kg; the known maximum weight is

'6For the Indian С. 1. pallipes, Pocock (1941) recorded a maximum body length of
100 cm and greatest weight of 24 kg (53 pounds). Weight is, evidently, sometimes more.

'7Since the complete list of these forms is of limited interest to our readers, it is
not given here.

The area of all North America is greater than the territory of USSR by only a little
more than 2 million km’.

198

175 pounds—79.3 kg; but on average their weight is less, and 45.4 kg
(100 pounds) is the weight of a large animal. The smallest wolves live
in the extreme south. Those of Texas and Mexico, С. /. baileyi, and
still smaller are those inhabiting the extreme southeast, С. /. rufus.
Weights of the males of this form reach 32—36.5 kg (70-80 pounds),
but on average the animal weighs 14—16 kg (30-35 pounds). In this
way, the American wolves follow Bergman’s rule more clearly and
more sharply than do the wolves of the Old World.

Coloration of American wolves also changes geographically in a
regular manner. In the extreme north and northwest live the northern
tundra wolves. Some races (populations) have a nearly white color (C.
I. manningi, С. |. bernardi, С. 1. tundrarum). Wolves of the forest zone
have a more saturated coloration, which generally corresponds to the
coloration of our forest wolves. Steppe, and especially desert forms of
the south and southwest have light ocherous tones. In the extreme
southeast of the range, in mesic portions of the country, in the area of
the lower course of the Mississippi and in Florida, the wolves have a
very intense color with well-developed ocherous and red tones (“red
wolf” of the Americans). Here, melanists are encountered with greater
frequency than in any other place, and the coloration of local animals
is generally dimorphic'®. In this way, Gloger’s rule is sufficiently clearly
revealed in the described color distribution (V.H.).

Biology”

Population. Within the boundaries of their range, wolves are very
unevenly distributed. Their distribution over a territory 1s conditioned
by the abundance and the stability of food resources, primarily wild and
domestic ungulates, and by the possibilities of capturing them, which
depends in winter on the depth and friability of the snow cover.

'SAmerican authors usually consider the wolves of the southeast as a separate
species, C. niger Bartram, with three subspecies (niger, gregoryi, rufus). This fact is
hardly well founded. Through the small С. /. /ycaon of the southwest, both groups of
races, “С. lupus” and “С. niger,’ merge with each other. Here, the forms of the niger
group are assigned to С. lupus.

'Information on the life habits, distribution and abundance of wolves appears in
a large number of journal notes and articles, and separate books, and in faunistic studies
and reviews, but only a part of this material represents scientific publications based on
original and accurately collected data. Therefore, the ecology of wolf cannot be consid-
ered well studied. Information on the biology of individual populations and subspecies
is especially scarce.

147

199

On the Kola Peninsula, in the second half of the past century, the
wolf was very rare, resulting from a decrease in the number of wild
reindeers and the reduction of domestic reindeer herds of the Saami
(Laplanders). Wolves at that time were relatively numerous along the
boundaries with Sweden and Norway, in their eastern regions where
the wild reindeer was preserved and where domestic reindeer breeding
was in better conditions (Pleske, 1887). In the Lapland preserve, in
1929—1938, wolves were totally absent and it was only later that they
began to bother herds of wild reindeer. In the winter of 1940/41, only
three wolves lived in the preserve in an area of 1300 kg? (Semenov-
Tyan-Shanskii, 1948).

In the Karelian ASSR, they [wolves] continually inhabited the
southernmost regions settled by humans. In certain years, wolves were
numerous in the Pri-Onega and Olonetsk regions, in Trans-Onega and
in those regions of the republic with little forest, especially along the
White Sea coast (Stroganov, 1949). The wolf is rare in the Segezhsk
region, and in the northern regions of the Karelian ASSR with deep
snow (White Sea, Kemsk and Loukhsk regions), it does not live con-
tinuously, appearing periodically every 5—10 years. Their appearance is
connected with the driving of domestic-reindeer herds or the migration
of wild reindeer (Marvin, 1951, 1959).

In the tundra and forest-tundra of the European part of the USSR,
wolves are relatively numerous; they accomplish seasonal migrations,
in summer going onto the tundra and in winter, into the forest-tundra.
In the Mezensk region, wolves are few in summer and numerous in
winter, since some of the migrants from the Kanin tundra approach
localities of resident wolves. In summer, only a few pairs remain to
raise their young on the Koidinsk tundra (Abramovsk coast of Mezensk
Gulf) and along the northern tributaries of the Nes’. On the Kanin
peninsula, wolves are encountered the year round. By the beginning of
winter, the majority of wolves wander into the forest-tundra; some
migrate to the Mezensk region, but the main mass winters on the
southern coast of Cheshsk Gulf. To this region also come wolves from
the Timansk tundra. In summer, not more than 3—4 pairs of wolves
remain in the huge area of the Cheshsk Gulf from Nes’ village to the
lower Pesh (along the Vitasu, Ome, Snope rivers).

On the Timansk and Malozemel’sk tundras, wolves den along the
Volong, Travyank, Shchuch’ei, Indige, Belaya, Svetlaya, Kamennaya
Visk, Vel’t’, Nerut, and Sul rivers and their tributaries. On the Timansk
and Malozemel’sk tundras, wolves remain in summer in the Timansk
range; near the coast, they are exceptionally rare.

200

On the Bol’shezemel’sk tundra no wolves are found in the repro-
ductive period in a coastal strip about 100 km wide, where there are
few suitable den sites, and in summer, herds of domestic deers are
pastured along with numerous dogs. Wolves den in the basins of the
upper courses of the Shapkinaya, Kuk, Lai and Kolva rivers (western
part of Bol’shezemel’sk tundra), along the upper Adz’va, Bol’shaya
Rogovaya and Chernaya (central part) rivers, and in the Pai-Khoi range
and the upper Korotaika, Silovaya and Kara (eastern part) rivers. In
winter, the majority of wolves migrate from the Bol’shezemel’sk tundra
to the forest-tundra, from the bend of the Pechora to the upper Usa;
some move beyond the Urals (Semenov, 1954).

Wolves are generally fewer on the European tundras than in
Arkhangel’sk taiga. On the Timansk tundra only 10 wolves were found
in an area of 10 thousand km’, or 1 wolf per 1000 km. In autumn, in
the event of successful reproduction, the number here could be 2.8
wolves per 1000 km? (Semenov, 1954).

In the taiga forests of Arkhangel’sk district, wolves are common in
the southern and central regions—in the basins of the Northern Dvina
and Onega. Along the valleys of these rivers, and sometimes of the
Pinega, wolves, singly and in groups, traverse northward. In the
Priozernii region, from 1947—1952, the number of wolves did not ex-
ceed 40—45 individuals in autumn (3 wolves per 1000 km’; Semenov,
1954). In Arkhangel’sk district, the wolf prefers places where the taiga
is thinned by cutting, the forests are intersected by roads and close to
villages where there are large areas of fields, floodplains and forest
cuttings since, in winter, the wolf finds its food only near humans. In
other northern districts with developing agriculture, the number of wolves
also increases, and the districts inhabited by them spread northward
(Semenov, 1954).

In the Komi ASSR, wolves are rare. In the taiga, they are only
encountered in the river valleys and lake depressions. They are more
common in the agricultural landscapes of the southern subzone of taiga,
and on the tundra. During 12.5 years (1939—1950) in the Komi ASSR
only 343 wolf skins were tanned. The average number tanned yearly
from 100 km? of taiga ranges from 0 to 0.01; in regions with agricultural
and other open lands, from 0.01 to 0.06 skins (Ostroumov, 1953).

In the Urals, wolves are most numerous in tundra, forest-steppe
and steppe. Their distribution there is also closely related to human
settlement. Deep in the taiga, wolves are few or totally absent (Shvarts,
Pavlinin and Danilov, 1951).

148

201

In Yamalo-Nenetsk Natl. Region, the wolf is met with almost
everywhere, but dens mainly in the southern tundra belt, in forest-
tundra, and in the thin forest of the pri-tundra. In winter, it remains in
the immediate neighborhood of domestic reindeer herds and 1$, there-
fore, very rare in the Arctic tundra. In the taiga, it only rarely appears
in the Krasnosel’kupsk region and in the southern parts of the Purovsk,
Nadymsk and Shuryshkarsk regions. During a decade (1948—1957),
1166 wolf skins were tanned from this area (85 to 157 per year)
(Rakhmanin, 1959).

In western Siberia, the number of wolves is high in the southern
tundra, in forest-tundra, southern taiga, birch forest-steppe, and steppe.
The northern and middle parts of taiga zone are distinguished by low

numbers of wolves to the west (the regions to the west of the lower and

middle courses of the Irtysh and the lower Ob’). Wolves only traverse
the region between the lower Irtysh and middle Ob’ in some years.
Thus, in the Surgutsk region, only 2 wolves in all were caught in 20
years, and in Vasyugan’, only two intrusions were recorded over 12
years. Breeding by wolves has never been recorded in these regions.

The somewhat unexpected northeastern maximum in wolf numbers
in western Siberia originates in the area with maximal depth of snow
cover, but this area is notable for its compact snow cover and the
abundance of wild and domestic reindeer. In the south, regions with
high wolf density coincide with the places of high number of roe deer,
regions of developed stock-breeding, and uneven snow cover (Laptev,
1958).

Along the Yenisei, in the taiga zone, the wolf is rare everywhere
until Turukhansk. In the forest-tundra (north of 76° N. Lat.), wolf
numbers noticeably increase. Places of wolf concentration on the tun-
dras are restricted to the regions of pastures of domestic reindeer herds
or stopping places of wild ones (S. Naumov, 1931). Wolves are com-
mon on the Yenisei delta; periodically they are very abundant along the
lower Olenek, Yana and Lena [rivers]. In central and southern Yakutiya,
wolves are few, but in Verkhoyansk territory there are more, and in the
Kolyma and Chukotsk lands, they are relatively numerous. Along the
Anadyr, they are common (Ognev, 1926; Sikol’nikov, 1928). In the
Lyakhovsk islands the wolf, being closely associated with tundra and
reindeer, is encountered only in summer and in winter migrates away,
following the deer herds to the mainland (Tugarinov, Smirnov and
Ivanov, 1934).

202

Fig. 52. Distribution and relative numbers of wolves in western Siberia (from
Laptev, 1958): 1—individual occurrences, 2—direction of migrations, 3—area of
migrations, 4—low population density, 5—high population density.

In the taiga zone of middle Siberia, in the Nizhnaya and
Podkamennaya Tunguska basins, wolves are most numerous in the
upper Khatanga-Kotui, Vilyuyi, Moiero and the northern tributaries of
the Nizhnaya Tunguska, where large herds of domestic reindeer graze.
Along the Podkamennaya Tunguska, to the lower course and southern
tributaries of the Nizhnaya Tunguska, wolves are scarce because of
deep, porous snow cover and the small number of wild and domestic
ungulates. Wolves usually appear here only for a short time (N. Naumov,
1934). This area of low numbers and nonpermanent habitation of wolves

149 on the right bank of the Yenisei almost joins with a like minimum in
wolf numbers in western Siberia (interfluves of the Ob’, Taz and
Yenisei rivers).

In Lena-Khatanga district, wolves are common along the Olenek

and along the Muna—a tributary of the Lena. In the Vilyuyi basin the

203

Fig. 53. Distribution and relative numbers of wolves in eastern Siberia (from
Khudyakov, 1957): 1—wolves numerous, 2—common, 3—rare, 4—1ransient,
5—wolves absent.

wolf is rare, and its numbers increase only. above the Arctic Circle
(along the Olenek). But in the valley of the Vilyuyi itself, they are
relatively numerous. In central and southern Yakutiya, and also in the
Kolymo-Indigirsk territory, the wolf is rare even in settled regions, or
is only found sporadically, but it is common on the tundra and forest-
tundra.

In the taiga zone of eastern Siberia, in the Sayans and Altai, the
wolf is scarce due to deep snow. These predators appear here only in
those regions where domestic or wild ungulates are common and where
thinning or reduction* of forests by humans was promoted by reduction
in porosity and depth of snow cover. In the Atlai and Sayans, wolves
are few, and in the majority of regions they only appear accidentally.
In the mountains, in autumn, they follow roe deer into the valleys,
where the snow is shallower and firmer (Polyakov, 1866; Solov’ev
1921; Afanas’ev et al., 1953).

In eastern Siberia, wolves are very abundant along the border with
the Mongolian Republic; in the southern Pri-Baikal, near Irkutsk, they

*Misprint in original Russian; svedenie (information) instead of snizhenie—Sci. Ed.

150

204

are common in regions along the railway (Pri-Baikal, Trans-Baikaliya).
The predators are rare or absent in the northern and a bit of the
southern taiga in regions of deep snow (Table 11).

In the taiga of the Far East, the wolf is also rare in all regions of
deep snow, especially those little opened up by humans. Wolves are
common in places with abundant wild or domestic ungulates and in
regions where the taiga has been considerably thinned by logging. In
Primor’e, the predator is rare—along the Iman (Ussuri territory), they
are encountered principally in its lower course (Zolotarev, 1936).

In the middle and southern zones of the European part of the
USSR, including the Pri-Baltic, wolves are rare. Their number grew
during war time, 1941—1945, but as a result of intensive control, at the
present time their numbers are again reduced. In Belorussia, the wolf
is met with everywhere. In 1932—1939 it was most numerous in Minsk
district, and later in Gomel’sk, Mogilevsk and Vitebsk districts (Table
12).

In the central chernozem districts, not long ago wolves were quite
common. Relatively large numbers were noted in 1951—1953 in
Voronezh and Belgorodsk districts (Barabash-Nikiforov, 1957).

In southern Bashkirtya at the edge of the upland-forest landscape
in the forest-steppe, wolves were few in the 30’s. About 1950, they
were quite often encountered, but in the upland broad-leaved forests,
they became scarce in winter (Kirikov, 1952). Wolf numbers here are
sharply reduced. In Volzhsk-Kamsk territory, from 1950 to 1965 wolf
numbers sharply decreased. In Tatariya during the five years from
1960—1964 the average annual production of skins was 170 (140—
186, V.A. Popov).

In the Ukraine, the wolves multiplied during the war years, 1941—
1945. In 1947—1949 their numbers in the entire Ukraine reached
about 7000. Population density was low in those places where vil-
lages were closely situated (Korneev, 1950). After 1945, wolves were

_ frequent in the southern part of Sumsk district (especially in more

wooded regions). In Chernigovsk district, which is very suitable for
wolf habitation, their high numbers were retained the entire time and
in 1938, 110 animals were caught here. After the war, especially
many appeared in Chernigov Poles’e; fewer in the forest-steppe and
southern regions. In Kievsk district, the predators became numerous
in its northern, forested sections. They were occasionally captured in
the regions adjoining the Poltavsk and Chernigovsk districts; in Kievsk
district, wolves were noted in a series of southern regions. In

151

205

Table 11. Manufacture of wolf skins from eastern Siberia (Khudyakov, 1937)

Years 1932 1933 1934 1935 Average,

Regions %
Main Western 398 315 309 292 11.0
Eastern 1478 1175 905 1369 42.4
Remote taigas 16 7 20 17 0.6
Northern 38 33 46 89 1377

Buryat ASSR 1636 1186 1266 1239 44.3

Table 12. Average annual output of wolf skins per 1000 hectares in
Belorussia (Serzhanin, 1955, with modifications)

Years, data 1932—1939 1948—1952
Average Fluctuation Average
by district in various

regions

Districts

Brestsk es — 0.5
Vitebsk 0.1 0.8—0.02 0.3
Grodnensk — — 0.5
Gomel’sk 0.2 0.80.03 0.3
Minsk 0.3 0.8—0.03 0.4
Mogilevsk 0.1 0.3—0.03 0.4
Molodechnensk — — 0.4

Zhitomirsk district, wolves were restricted to the northern regions,
along the border with Belorussia (Korneev, 1950).

In the western districts of the Ukraine, wolf numbers are generally
not great. They are higher in western Poles’e, in Rovensk and Volynsk
districts. There are many wolves in the montane regions of former
Stanislavsk district. In the Trans-Carpathians and in L’vovsk district,
the wolf is rare; in Ternopol’sk district, it does not live permanently,
appearing periodically from the southern regions of Rovensk and the
northern regions of Stanislavsk districts (Tatarinov, 1956). In L’vovsk
district, wolf litters were noted only in the Kamensko-Bugsk, Sokal’sk
and Peremyshlyansk regions. In the Trans-Carpathian area, they are
relatively common in all montane and low-lying regions (except rocky
places in the high mountains). Stray wolves were met with in the
Vinogradovsk, Uzhgorodsk, Perechinsk and Mukachevsk regions. In
Drogobychsk district, wolves are common in montane regions; in 1949
litters were encountered in the Turkovsk and Podbuzsk regions, and
strays in the Strelkovsk and Dorogobyshsk regions.

206

In Stanislavsk district, wolves occupy permanently the southern
and southwestern montane region, and intrude on the plains. In
Chernovitsk district, wolves occur permanently in five southwestern
regions bordering Romania, and also in Sakyrsk and Kal’menetsk re-
gions, where they often come from Moldavia.

In Kamenets-Podol’sk district, wolves are established in the north-
ern regions, but animals are scarce here. In Vinnitsk district, wolves
are regularly noted only in two southern regions; they penetrate hither
from Moldavia, where there are many wolves. In Kirovogradsk district
wolves are more or less constantly present in the Pri-Dneprovsk re-
gions adjacent to Kiev district. In Poltavsk district, wolves live in the
eastern regions (near Sumsk district), and penetrate along the forested
massif into Mirgorodsk region; they were also found nearer to Kiev
district.

In Khar’kov district after the war, wolves propagated more in the
central regions. Very few litters were found in the Volchansk region,
and in the Kupyansk region vagrant wolves were recorded. In the
Lugansk district, wolves were present in 31 out of 34 regions; they
were especially numerous in the northeastern regions. In six of these,
in 1949, 150 wolves were captured. In Donetsk district, wolves are rare
and noted only in the south. In Dnepropetrovsk district after war, wolves
lived permanently only in the extreme eastern regions. In Zaporozhsk
district, wolves live mainly in places near the Dnepr floodlands, and
appear occasionally in the Berdyansk and Pri-Azovsk regions.

In Khersonsk district until 1948—1949, the Tsyurupinsk and
Skadovsk regions were densely inhabited by wolves; on the right bank
they were totally absent; after 1948, they appeared in the Khersonsk,
Berislavsk and Novovorontsovsk regions. In Nikolaevsk district, wolves
live permanently in the Shirokolanovsk region; strays appear in the
western regions. In Odessa district, wolves are common in the western
regions. In 1949—1950, strays were also observed in the central re-
gions. In the Izmail’sk region, wolves live permanently in the regions
adjacent to the Moldavian SSR. In the floodlands of the Dnestr, wolves
are few (Korneev, 1950). In the Crimea, wolves were exterminated, but
during the years of the Patriotic [2nd World] War, they appeared again
on the peninsula; they were soon destroyed.

In the Caucasus, the wolf is distributed everywhere, though not
evenly. According to the data on tanned skins, the greatest number of
individuals per unit area was captured in the western Cis-Caucasus and
eastern Trans-Caucasus. In the 1930’s of this century in Azerbaidzhan,

152

207

were caught per 1000 km’, 11.6 wolves; in Armenia—9.9, in the north-
ern Caucasus, 6.7; in Dagestan, 6.7; and in Georgia, 0.8 wolves. The
lower Terek, Sulak, Kuma and the coast of Kyzylagachsk Gulf were
the least densely populated by the wolf. In alder bogs of Kokhida, wolf
is common in the lower Gagid, Okum and Picher rivers. In montane
regions of the Caucasus, wolf numbers are high in the Caucasian,
Borzhomsk, Zakatal’sk and Lagodekhsk preserves (Vereshchagin, 1959).
In the Trans-Caucasus, wolves are everywhere, but along the wooded
Black Sea coast they are so rare that they were never seen by the
majority of inhabitants (Satunin, 1915).

In Kazakhstan, the wolf is encountered nearly everywhere. Their
numbers are highest in the west, in several regions of Aktyubinsk and
Kustanaisk districts, in the valleys of the Syr-Dar’ya, Chu, and Ili
rivers, and in the mountains and foothills of the Zailiisk and Dzhungarsk
Alatau, near Lake Kurgal’dzhin, in the Alakul’sk depression, around
Zaisan Lake and in several other places. In the northern forest-steppe
regions, wolves are scarcer. However, in the former Kustanaisk district
of the former Turgaisk governance, their number in beginning of the
twenties was extremely high (see beyond, V.G. Heptner). In clay deserts
and semi-deserts, where there is neither water, wild ungulates or live-
stock, these predators are absent or appear there only from time to time.
There are also no wolves in some of the montane taiga regions of the
southern Altai, for example, in the Zyryanovsk region, where snow
cover is deep and porous (Afanas’ev et al., 1953). The general number
of wolves in Kazakhstan is estimated at approximately 25—30 thou-
sands. About 10 thousand wolves are exterminated annually in
Kazakhstan; i.e. a little more than one third of the population (Afanas’ev,
1955).

In Kirgiziya, wolves are encountered everywhere. They are most
numerous in the regions of large high montane pastures-elevated
watersheds, since in these places not only many livestock are found but
also great marmot colonies were preserved, and in the neighboring
ranges, large numbers of wild ungulates (arkhar sheep and wild goats).
Each year in Kirgiziya, prior to the Great Patriotic War, up to 400—
600 wolf skins were tanned (Kuznetsov, 1948).

In Uzbekistan, there are few wolves. In Kashkadar’insk district
they are rare; they are rarely met with in desert regions, even in winter;
in montane regions, they are numerous (Meklenburtsev, 1958).

In the Turkmen SSR, the distribution of wolves is restricted by the
presence of water sources and the abundance and also availability of

[53

208

Jot]

=a)

Fig. 54. Relative number of wolves in various parts of Kazakhstan (from
Afanas’ev et al., 1953): 1—general distribution, 2—districts where wolves are
especially numerous.

food. In Badkhyz (southern Turkmeniya), there are more wolves, ap-
parently, than in other regions of Turkmentya, which is explained by
the abundance of wild and domestic ungulates. The especially large
number of wolves between the Tedzhen and Murgab has already been
shown by Radde and Walter (1889). Along the Kushka river wolves
frequently hunt near the most remote villages. Along the Egrigek river,
there are more wolves than along the Kushka; although here there are
no human settlements and livestock, dzheiran gazelles [Gazella
subgutterosa] are numerous, which come to the river for drinking and
there are also abundant watering sites and convenient places for dens
and diurnal lairs. In Gyaz’-Gyadyk, wolves are common in the whole
region. There are many wolves along the Tedzhen river. In the desert
part of Badkhyz, the wolf is everywhere, but is unevenly distributed,
being concentrated by wells, in grazing sites of sheep flocks and near
waterholes, especially those situated not closer than 100—200 m from
domiciles (earthen huts). Six to seven wolves on average are associated
with one waterhole. In winter, wolves are evenly distributed in Badkhyz,
because of the reduced need to drink and the wider pasturing of the
sheep flocks (Heptner, 1956). The number of wolves in western
Turkmeniya is high, but there distribution is uneven.

It is difficult to estimate, even approximately, the general number
of wolves in USSR, not only because of the absence of data, but also

209

because of uninterrupted changes in their numbers. In the 1930’s, their
number in our country was estimated by a series of authors to be 60—
80 thousands, a figure which was, apparently, an underestimate. At
nearly the same time, S.A. Buturlin evaluated the wolf population as
being not less than 100,000 individuals, and the yearly production as
47,000. After the Patriotic War of 1941—1945, the number of wolves
markedly increased, particularly in the regions which suffered from the
German invasion. After restoration of the destruction caused thereby,
as a result of augmented wolf control and the application of a series of
very effective means for their extermination (shooting from aeroplanes
and airosled*, etc.), the number of wolves was reduced in the 1960’s
by not less than 3—5 times and in places, more. In a series of districts
they disappeared entirely.

One may establish several general laws:

1) The highest numbers of wolves are in so-called “open” land-
scapes—southern tundra, forest-tundra, forest-steppe, steppe and several
desert regions.

2) Within these zones, the distribution of the predators and their
mode of life, in particular nomadism, depends upon the allocation of the
chief foods—wild and domestic ungulates.

3) In the forest zone, especially in the taiga, the most important
obstacle for wolves reveals itself to be deep, porous and prolonged
snow cover, which sharply limits the possibility of movement and cap-
turing food. The rarity or absence of wolves in the deep taiga regions,
which is a consequence of this situation, was already established by
A.F. Middendorf (1869)** and later confirmed by all investigators.
However, a clear coincidence between the snow cover depth and the
abundance of wolves cannot be established, since the relation here is
more complicated; in actuality, the influence of snow on the number of
wolves acts through a complicated system of biological relationships
and essentially changes through the direct and indirect effects of human
activities.

The construction of roads and paths increases the possibility of wolf
movements, while the added complexity of the landscape and the appear-
ance of farming and animal husbandry in these regions enriches the potential
food reserves, especially in winter time. This explains the direct relation
that is manifested between the number of wolves and the agricultural
development of the territory in little-developed taiga regions.

*Forerunner of snowmobile—Sci. Ed.
**Not in Lit. Cit—Sci. Ed.

154

210

In many poorly inhabited regions of taiga, there are no permanent
wolf populations and the animals only rarely intrude into these places
by chance. In such unfavorable and almost uninhabitable territories,
wolves penetrate mainly along the river valleys with their rich and
complex biocenoses and their greater degree of development of places
suitable for humans.

4) In the desert-steppe regions, the presence of watering places
serves as the important factor limiting the number and distribution of
wolves, being quite necessary for the predators, especially in the hot
time of the year. There, concentration of the animals is noticeable near
wells and in river valleys, where they find, not only the necessary
water, but also abundant food since in such places both wild and
domestic hoofed animals are concentrated.

Habitat. Places of diurnal rest, where dens are also located during
the reproductive season, are usually especially well protected and are
characteristically near watering places. The hunting territories of wolves
are various, and are restricted only to the possibilities of capturing food.

On the tundra, places of diurnal rest of wolves are restricted mainly
to river valleys and thickets in dry plateau sections; in the forest-tundra,
forest clearings and osier-beds and thickets of dwarf arctic birch serve
this purpose. In the Yamal and Bol’shezemel’sk tundras, wolves con-
struct dens, mainly among bushes in valleys and more rarely in the dry
meadows of slopes; they use flood terraces and dry osier-bed thickets
along drainage divides. Of 11 biotopes in river valleys, wolves use
three, and on interfluves of dry plateaus, of 17 biotopes, they are
distributed in only one (dry shrubs along the divides; Sdobnikov, 1937).
Wolves colonize sea coasts only residually*, where they often feed on
marine flotsam (Sokolov, 1951).

In the forest zone, wolves avoid monotonous dense forests. In the
taiga of West Siberia, they live chiefly in the river valleys (floodplains)
(Laptev, 1958). In the Irkutsk district, they prefer places where small
sections of forest alternate with fields, and in winter they use roads and
paths laid out by humans widely (Timofeev, 1941). In the Karelian
ASSR, wolves live mainly near populated places, in cut-over forests
and shrubs adjoining fields. In winter time, they not only approach
populated places, but often enter them (Marvin, 1959). In Byelorussia,
avoiding the large forest massifs, they prefer small dense plantations
with thickets (Serzhanin, 1955).

*The Russian word gushcha (sediment, dregs) is used here—Sci. Ed.

155

211

In heavily settled regions, wolves, though living in the immediate
neighborhood of villages, stay in especially secluded places: forest
islands, bogs, remote ravines and gorges, and dense growths of shrubs;
at the same time, they often live in vacant weedy field, and sometimes
in crop fields (Mertts, 1953).

In the steppes, wolves spend the day, and in the reproduction
season construct dens, in growths of shrubs and the remote ravines and
gorges overgrown by weeds, among tugais [riparian thickets] in the
valleys of rivers and streams, in the old weedy fallow fields and in crop
fields. In western Kazakhstan, wolves are common in summer in reeds
by water bodies, in ravines and sand hills, if there are waterholes there.
In winter, they concentrate near livestock pastures, in reeds along the
banks and by large lakes, and also near villages, where they feed on
carrion and hunt dogs. In northern Kazakhstan, wolves live in narrow
gorges, in vacant lands among weeds, and in forest clearings choosing
elevated, drier places. In winter they spend the night there and gener-
ally remain close to human settlements which they approach by night,
while during the day they hide in weeds or reeds by lakes (Afanas’ev
et al., 1953). In the Kustanaisk intact steppes, they find shelter in
growths of dwarf almond (Amygdalus nana) and steppe cherry (V.G.
Heptner).

In the deserts and.semi-deserts of central Kazakhstan, wolves are
restricted in the warm time of year to melkosopochniks near springs
and streams, in deep sayakhs* (ravines and valleys) if water sources
are found in their bottoms, in weeds by water bodies and in forest
islands. In winter, they concentrate in regions of livestock husbandry,
near rivers and lakes, or they follow the herds of saigas and dzheiran
[Gazella subgutterosa] (in Betpak-Dala).

In southern Kazakhstan, especially numerous are wolves living in
the river valleys (Aksu, Karatal, Ш, Chu, Talas, Syr-Dar’ya). In sum-
mer, they are found in the tugais, reeds and rose willows at the lake
channels, and rarely in the stabilized sand hills near water. Here, many
wild pigs, roe deer, hares, pheasants and waterfowl are found year-
round; here also domestic cattle graze. In winter, the wolves move, by
night, from the river valleys to adjacent regions where domestic cattle
and dzheiran winter, but at daybreak they again return to the valleys.
In the sands, wolves only occur near waterholes, constructing dens in
saxaul forests or in dense growths of shrubs (Afanas’ev et al., 1953).

*Foreign (Turkic?) word—Sci. Ed.

212

The distribution of wolves among the biotopes in other desert re-
gions of Middle Asia is the same. In Turkmeniya, the wolf is widely
distributed, but is obviously attracted to human settlements and places
where domestic livestock graze. It is encountered high in the mountains
(2000 m above sea level and higher), in sandy and clayey deserts and
cultivated landscapes. In intense heat, wolves are almost absent deep
in the desert, or they are rare and are restricted to only the few wells
which are used by people where there are sheep flocks and also water
remaining from the drinking places of the herds. The animals occur
mainly along the river valleys near the watering places of wild ungu-
lates (mainly dzheiran) and domestic cattle. In winter, when the wild
ungulates and herds are distributed throughout the whole desert, wolf
distribution changes accordingly (V.G. Нершег).

In the mountains, the wolf ascends to a height of 3 and 4 thousand
m above sea level (Pamir). In the mountains, the thick and very snowy
cover favors wolves because they, in following ungulates, stick to less
snowy slopes and those blown free.

In the northern Caucasus, wolves prefer inhabited places, and in
remote places, are met with only where there are many wild ungulates.
The life of the wolf here is closely connected to the domestic animal
herds, especially sheep flocks, in the following of which the predators
ascend the mountains in summer and in winter, descend (Satunin, 1915).
In the Caucasian preserve, 30% of the incidence of wolf encounters
were in the alpine zone, in dark coniferous forests, 32%, and in the
broad-leaved forests, 38% (Teplov, 1938).

In Armenia, the wolf inhabits the dry subtropics, semidesert, mon-
tane and meadow steppes, sub-alpine and alpine meadows, and also
encountered in barren mountains, cliffs and talus, and do not avoid
cultivated lands. Within the boundaries of Armenia, the wolf is met
with at heights of 560-3800 т above sea level. In summer, a greater
portion of the wolves hold to high mountains, in places where domestic
cattle graze, and are in small number in the desert and barren mountain
zones. By winter, wolves descend to the valleys and only a few of them
remain in the mountains near herds of wild ungulates (Dal’, 1954).

In the mountains of Kazakhstan and Kirgiziya, wolves remain in
summer in alpine meadows, abounding in marmots and wild ungulates,
and where the domestic animals pasture at this time. In the upper
reaches of montane rivers (at heights of 3000—3500 т above sea level)
wolves also raise their cubs at this time. After the development of snow
cover in the mountains, following the wild and domestic ungulates, they

156

213

descend to the foothills and valleys; only a smali number of them
remain near the herds of wild, and sometimes domestic, ungulates,
remaining in the mountains in winter in the heat of the sun in sunny and
wind-blown areas (Afanas’ev et al., 1953).

In Semirech’e there are, apparently, two populations of wolves;
one lives in the mountains and descends in winter together with the
ungulates, wintering in the foothills, while the other—the plains popu-
lation—occupies in summer impassable thickets of sea buckthorn,
tamarisk, and other shrubs in the tugais. Both populations are associ-
ated with flocks of sheep and wild ungulates (Shnitnikov, 1936).

Food. The wolf is a predator, the main food of which consists of
mammals of moderate and large size. Almost everywhere it is associ-
ated with the ungulates comprising its main food. The quantity and
accessibility of it determines the predators’ numbers. Upon the life
characteristics of the ungulates depends also the way of life of wolves,
differing in different regions because, first of all, the species composi-
tion and biology of the wild ungulates or the methods of keeping domestic
animals in these regions are not the same. If not fundamental, then of
significant importance in the nutrition of wolves are animals of medium
and small size—marmots, hares, badgers, foxes, polecats and others. In
many places, wolves successfully hunt dogs, and, in the regions where
raccoon dogs are acclimatized, they are destroyed in great numbers by
them. Among small mammals, wolves catch ground squirrels, mice,
hamsters, voles and other rodents, and also insectivores. In the agricul-
tural forest-steppe and steppe regions, wolves eat small rodents in the
fields. They successfully hunt waterfowl, especially during the time of
their molt. Large birds, mainly egg clutches and young, suffer from
wolves. Wolves destroy many domestic and wild geese. Reptiles (liz-
ards and snakes), frogs, rarely toads, and also large insects are eaten
by wolves when other foods are insufficient. In times of hunger the
predators readily eat carrion, visiting cattle burial grounds, slaughter-
houses, tallow-presses or especially situated lures. If such places of
carrion deposition are permanent, they can determine the winter route
of wolf packs.

In many, if not all, regions, wolves are partial to plant as well as
animal food. They willingly eat berries of mountain ash, lily of the
valley (Convallaria majalis), bilberries, blueberries and cowberry (in
the forest zone), nightshade (Solanum nigrum), fruits of apple, pear
and others (in the south). In summer, they readily visit melon fields, eat
watermelon, muskmelons and frequently cause great damage, not only

214

by eating the fruits, but many others are spoiled. They frequently eat
various grain crops, and in the Pri-Ural steppes, the delicate and sweet
shoots of reeds.

Cannibalism is not rare among wolves. In times of hunger in win-
ter, the pack often attacks weak or injured animals. They may attack
a male, which in its struggle for a female, becomes severely wounded.
In captivity, cannibalism is noted during the transition of young wolves
from meat to milk or vegetable foods. The stronger cubs attacked and
ate the weak (Barabash-Nikiforov, 1957). Hungry wolves fight fiercely
for food, and frequently kill the weaker ones, which are afterward
almost always consumed. Cases are described when wolves killed and
ate the wounded or the dead bodies of their kin (Makridin, 1959).
Therefore, these animals are generally not fastidious about food, but
when possible, they eat only the best food; this is particularly related
to the summer time and concerns plant foods, the need for which is not
so large.

The wolf is an animal of great endurance. It is able, without losing
strength and running speed, to survive prolonged hunger, for a week or
more. But upon succeeding in a hunt, it is very voracious and, accord-
ing to some authors, can eat immediately a large quantity of food—up
to 25 kg, as if laying in a store (Manteifel’ and Larin, 1949; Timofeev,
1949; Semenov, 1954). It was noticed that a litter of 7-10 wolves
completely consumed an entire horse carcass during one night (Sokolov,
1951). In the lower Ili river (Kazakhstan) a pair of wolves ate a roe
deer weighing 25—30 kg or a young wild swine of 30—40 kg at one time
(Afanas’ev et al., 1953). In Badkhyz (Turkmeniya) a wolf was ob-
served to devour a young arkhar sheep of about 10 kg in weight
(Heptner, 1956). However, these amounts do not characterize the quan-
tity of food actually eaten at one time. A part of this food is usually
pulled apart and hidden away, especially when there is plenty of food.
At the same time, animals torn apart by wolves are often further eaten
by hyaenas, jackals and especially vultures. In Badkhyz, a skeleton of
a donkey which was torn apart by a pair of wolves at the start of night,
was found in the morning completely cleaned of meat (Heptner, 1956).
More than 1.5—2 kg of food can be rarely found at one time in the
stomachs of wolves (Novikov, 1956). According to the accurate data
of P.A. Martts (1953), a wolf eats no more than 3 kg of food at one
sitting and any excess amounts are quickly regurgitated. The relatively
small amounts of meat eaten by a wolf at one time is also to be
understood by the figures given above on the weight of wolves (p. 174).

157

215

The foods of wolves vary substantively in the various seasons, and
these changes are associated with changes in the life habits of the
predators—their transition from a settled life in the warm time of year
to a nomadic way of life in winter. In summer, various foods are
available for wolves, and its quantity is maximal in this period. There-
fore, summer feeding of wolves is diverse. The principal share in various
regions is occupied by different foods, but mainly by animals of me-
dium and small size. The significance of ungulates is reduced in this
period, although wolves still hunt them. Thus, in the stomach of a
young wolf killed on 3 July 1944, in the Bashkir preserve, remains of
a young maral [Cervus elaphus], a mole’s foot and 2 nestling sparrows
were found; and in the stomach of a juvenile wolf caught on 17 August
1941, 16 lizards were found. In the valley of the Ural river, wolves in
summer frequently visit melon fields and eat watermelons, and in the
Shaitantau [mountains] during a year of steppe-cherry harvest, its stones
were found in the majority of the wolves’ feces (Kirikov, 1952). At the
end of summer and the beginning of autumn wolves hunt pastured
livestock and wild ungulates, catch hares, dig up muskrat burrows and
houses, catch the small animals and birds, and around water bodies,
successfully obtain molting waterfowl.

After snow accumulation, feeding worsens. At this time, the prin-
cipal attention of wolves is directed to ungulates. In the time of greatest
hunger, wolves approach populated areas, hunt dogs, attack livestock in
their sheds even by day, and willingly make use of cattle burial grounds
and carrion.

In winter, wolves travel along roads and unwillingly turn into the
snow upon the appearance of not only a single sledge but even a series.
At this time, wolves also attack elk*. However, the attack by an indi-
vidual wolf of an adult elk, especially in more or less deep snow, often
ends in the death of the wolf. For instance, in the winter of
1952/1953 in Verkhne-Toemsk region of Arkhangel’sk district, two
wolves were found killed by elks (Semenov, 1954). An attack by a
pack usually is successful. In March—April, in the forest zone on crusted
snow, wolves rarely appear near the villages and mainly travel through
the forests, hunting roe deer, elk and [rein]deer on the snow crust.

Early spring (after the crusted snow) is the time of greatest hunger,
when wolves are very harmful to animal husbandry (especially in the
steppe), exterminating mainly the young. Chiefly at this time in the

*In North America, called ‘“‘moose’”—Sci. Ed.

158

216

steppe and desert, but also in tundra, wolves hunt pregnant ungulates
(dzheiran, saiga, roe deer and [other] deer). At the time of birth of the
young, wolves gather near the place of calving, where they kill both
adults and young.

After melting of the snow and the beginning of spring reproduction
of animals (end of April-May) wolves transfer their feeding mainly to
vertebrates of moderate and small size. In June, the young begin to feed
mainly on animals, and the mother’s milk is only a secondary food for
them. In June, wolf-cubs already can walk independently to water-
holes. From August on, attacks of domestic livestock become common
(Khudyakov, 1937).

The food habits of wolves in different regions differ markedly. The
food of tundra wolves during the snowy period is without exception wild
and domestic [rein]deer (mainly calves and does). They attack hares,
arctic foxes and other animals. In the Nenetsk Nat. Okr., in the stomachs
of 74 wolves caught in winter and spring were found remains of: rein-
deer—93.1%, small rodents—S.4%, willow grouse—4.1%, white
hare—1.3%, foxes—1!.3% and fish—6.8%. Of great importance in the
life of wolves living at the expense of ungulates is the group attack and
mass “cutting” of the prey into [smaller] flocks and herds. Later, wolves
often travel to places of these “foragings” to utilize the preserved re-
mains. Small groups of predators remain near the sea coast or by villages
where they feed on marine flotsam along the shore, the wastes of indus-
try, carrion, and rob the traps and snares of hunters (Makridin, 1959).

In the summer food habits of wolves on the tundra, a considerable
role is played by birds (in the second half—especially molting geese
and ducks) and small rodents (lemmings and voles). [Rein]deer, espe-
cially at the beginning of summer (calving time), also play an important
role. In the montane regions in the northeast, snow sheep, hares and
marmots occupy a notable place in the food habits of wolves (A.
Romanov, 1941).

In the taiga of Karelia, wolves attack elk (young), reindeer and
domestic livestock and eat carrion; in summer, they catch rodents,
ground nesting birds, and sometimes lizards and frogs. They also eat
berries, in autumn especially mountain ash (Marvin, 1951).

In the forest of the Tatar ASSR wolves feed in the snowy period
mainly on mammals (98% of occurrences), especially the domestic
animals, and carrion (68%), followed by mouse-like rodents (24%)
and hares (21%). The proportion of birds constitutes only 10% of the
findings (as % of total number of stomachs, feces and food remains

159

217

investigated; У. Popov, 1952). In the region of the Rybinsk reservoir,
wolves in winter hunt mainly elk (Kaletskaya, 1953). In the Urals, they
feed on wild ungulates, hares, ground squirrels, domestic animals and
birds. In winter, they eat carrion and frequently pursue foxes (Shvarts,
Pavlinin and Danilov, 1951). In Belovezhsk Forest, they hunt chiefly
wild ungulates (48%), especially wild boars (21%), roe deer (18%),
[red]deer (6%). Remains of domestic animals were encountered in 28%
of the stomachs. They frequently catch brown hares (16%). In the
warm period of the year, the significance of ungulates decreases to
40% (in spring) and 31% (in autumn). At the same time, the role of
domestic animals in their food increases from 32% to 42% (sheep
particularly are caught). Wolves here hunt dogs relatively rarely (Gavrin
and Donaurov, 1954).

In the forest-steppe of the central chernozem regions, domestic
livestock, hares and small rodents comprise the principal food of wolves.

In steppe regions, in 56 cases (stomachs, food remnants) first place
was occupied by mouse-like rodents (35%), then carrion (17%—ainly
in the winter), dogs, calves, sheep, goats, swines (16%). In the stom-
achs, remnants of hares (8%) and ground squirrels (5%) were often
found; in one case, red fox. Proportion of birds (mainly domestic geese)
constituted 4%. Occasionally wolves eat lizards and insects (dung
beetles). Plant food are represented by berries, strawberries, pears and
apples (windfall). In the Usmansk forest, the usual prey of wolves
consists of deer, beavers, hares and raccoon dogs (Table 13). In sum-
mer and autumn, remains of dogs (38%), hares (18%) and sheep (13%)
are found in their food (Mertts, 1953).

Vertebrates are encountered in 92.2% of the cases in food habits
of Ukrainian wolves (Korneev, 1950).

Foods consist of mammals—90.7% occurrence, birds—12.9%,
reptiles—5.5%, amphibians—29.6%, fish—18.5%, insects—46.2%,
plants—48.1%. Among mammals, domestic animals are encountered in
48.9% of occurrences, and game species in 32.6%, including hares—
22.4%, roe deer—10.2%, mice—14.2%, voles—42.8%, shrews—6.1%.
Among domestic animals dogs occupy 18.2%.

Up to 10 water voles (Arvicola terrestris) and 15 common voles
(Microtus arvalis) were found in the stomach of one wolf. An espe-
cially great importance in the food of wolves is assumed by small
rodents in the years of their mass reproduction (Migulin, 1938). Among
birds, mallard (Anas platyrhyncha)*, domestic hens and geese

*Sic: should read platyrhychos—Sci. Ed.

158

218

Table 13. Food habits of wolves in the Usmansk forest (112 stomachs and 63
feces, Mertts, 1953)

Summer

Season, data Winter

Number of % Number of %
Food type occurrences occurrences
Deer 37 Dies 3 5.3
Beavers Di) 153 3 5.3
Hares 34 19.4 10 17.9
Mouse-like rodents 14 8.0 5 8.9
Raccoon dogs 7 4.0 1 1.8
Domestic dogs 20 11.4 21 37/5
Sheep 7 4.0 7 12.5
Swine 4 2.2 1 1.8
Domestic birds 4 DP 3 5.4
Carrion, bait wi 9.6 — —
Plant remains 5 2.8 2 3.6

predominate, among reptiles—lizards and among amphibians—toads.
Among fish, the pike (Esox /ucius) was found, being caught by wolves
in water-meadows (in times of flood). Other kinds of fish are caught
along the shore. Among insects, beetles (100% of occurrences),
Orthoptera (48%) and Hymenoptera (44%) predominated. Among the
plant food, berries of buckthorn (Rhamnus cathartica; up to 389 in one
stomach), black-berried nightshade (Solanum nigrum; up to 9082 seeds
in one stomach), berries of lily of the valley (Convallaria majalis; up
to 486 seeds in one stomach) and fruits of pear (Pirus communis; 140
seeds in one stomach) were met with (Komeev, 1950). In addition to
animal foods, many grains of maize were found in wolves’ stomachs
in the Caucasus, and in Kiev district, the fruiting bodies of mushrooms
of the genus 7richoloma (Komeev, 1950).

In the Caucasus preserve the frequency of occurrence of various
groups in the food of wolves was as follows (Teplov, 1938): mam-
mals—90%, ungulates—81%, wild boar—38%, deer—16%, tur—12%,
chamois—12%, roe deer—7%, rodents (hares and mouse-like)—90%,
carnivores (bear, fox, marten}—3%, birds (chiefly Caucasian black-
cock)—7%, fruits including berries (pear, apple sweet cherry, viburnum,
blackberry, dog rose)—12%.

In the Uralo-Embensk desert, the summer feces and food remnants
of wolves (268 examples) consisted of 37% mouse-like rodents, jer-
boas—9.1%, ground squirrels—13.6%, hares—2.6%, hedgehogs—2.9%,
domestic livestock—8.8%, birds—1 5.8%, reptiles—0.3%, insects—1.1%
and plant remains—5% (Kolosov, 1935).

160

219

In the northern zone of Kazakhstan the prey of wolves in summer
are small rodents (especially water voles), hares, young and molting
ducks, young blackcocks and willow grouse, and rarely roe deer and
domestic livestock (sheep). During times of drying-out of lakes, water
birds and water voles are especially available to wolves and hence
become the principal food, on which they also feed their litters of
whelps. In Kazakhstan the wolves also readily eat the fallen apples and
pears, and visit melon-fields. In winter in northern Kazakhstan, wolves
hunt domestic livestock, roe deer and eat carrion (Afanas’ev et al.,
1953). Around Lake Kurgaldzhin, wolves live all year round in the
reeds. In summer they feed on water voles and waterfowl, especially
during the time of molt. In winter, they live at the expense of wild pigs,
following them in paths which the swine tread in the snow.

In the Betpak-Dala desert, wolves feed in summer on dzheiran,
saigas and hares; they also eat gerbils, jerboas, turtles and insects. In
autumn and winter, the saigas and dzheiran which overwinter here, as
well as the domestic livestock suffer greatly from wolves. Mammals
predominate in wolves’ food in southern Pri-Balkhash (92—100% of
occurrences), among them ungulates (16—100%) and rodents (10-84%)
predominate. Among ungulates, wild swine (10-50%) and roe deer (S—
100%) are of greatest importance. The wolves frequently eat tolai hares
and muskrats. Around water bodies, they eat fish (mainly carp). Rarely
they attack foxes, and among birds, mainly waterfowl and pheasants
(Table 14).

In the lower Ili river the infrequent remains of domestic animals in
wolves’ food is explained here by the abundance of wild foods. Feces
containing soil indicate the necessity to wolves of melkozem [fine
earth] especially during the transition from summer foods to winter
ones and vice versa (Afanas’ev et al., 1953).

Domestic animals have important significance in the food habits of
Turkmenian wolves; as does the wild fauna, especially ungulates which
play a great role, particularly for the Badkhyz wolf population (south-
em Turkmeniya). This is, obviously, connected with the richness of
Badkhyz in wild ungulates. The particular attention of wolves here is
concentrated on dzheiran as the largest amount of the more available
species. Adult dzheiran are caught mainly at waterholes by wolves
concealing themselves behind dense thickets. Cases of attacks on kulans
by wolves are also known, especially during winter snowfalls and on
ice-covered ground. Small animals, especially reptiles, are known to be
significant in the food habits of wolves in the Badkhyz (Heptner, 1956).

————————eeEOeeeeeeeeee eens nn eS SSS
злеэА snoueA Ul

Pe = ee ss) VIG BAG (Ge (Gy) 0 — 0 бр 86 s9dUeLINDDO цвэш
ЗИ ВИ ОС Г ЕС ВЕС Ги 85 То. ЭГ 956025 SE-C 86 $5 001 8 I 0С 9 ES. Св: СЗ ul 540421351]

(% ul)
0 OL ГС 987 9711 BSS 8'0 70 ГЕТЕ OFF PPS OUALINDDO UPD

Га]
S
со
a)
<
д
<
+t
\o
—
ee
S
(99)
\o
д
+
>
+
S
(oe)

SUC ey
$19410
риа
x04
$19199
sjuapoy

pooj

sosn| [Oy
$199$и]|

Joop 901 Зипол
Jeog рим Buno x
1ээр 901 упру
тео4 рим уару
soyejn3sun
рэо3-иэлЯ

ЗЭ[ол 1238 М
$12115 ПИ

зэлеч во 1,

95998 рие syonq
sjelulue 215эщ0а

(SUOTJBIIFIPOU YIM ‘6961 ‘в 39 Aa SBuBJY) зэ4шехо Jo лэдшпи [6303 Jo зиээлэ4 SE OIG
—sojduiexa Jo лэашпи ‘661-6561 ромчэ4 эцз и! иэзеэ POO} рив SadaJ Jo SISA[EUY ‘дэлы ИТ доме] э4з UT зэлюм Jo $14еч роод “py эает 1910‘

—

221

Wild ungulates (goats, arkhar sheep, roe deer, maral) and domestic
livestock serve as the main food of montane wolves in Kirgiziya. Wolves
follow them into the high mountains in summer, and in winter, they
descend to valleys. In summer, wolves readily and successfully hunt
marmots, concealing themselves and waylaying them; they eat also
small rodents, birds and carrion (Kuznetsov, 1948). In the Dzhungarsk
and Zailiisk Alatau, the marmots frequently serve as the main food of
wolves (Afanas’ev et al., 1953).

Change in the quantity of the chief foods or the conditions under
which it is obtaining in different years strongly changes the character
of wolves’ food habits. In winters with much snow and particularly
with severe and protracted snowcrust, wolves sometimes almost com-
pletely exterminate the wild ungulates (especially roe deer), even over
great areas. Thus, during the winter of deep snow in 1940/1941 in
northern Kazakhstan, wolves almost exterminated roe deer in Kustanaisk,
North-Kazakhstansk, Pavlodarsk, Kokchetavsk, Akmolinsk, and East-
Kazakhstansk districts entirely. In the Presnogor’kovsk region of
Kustanaisk district alone, remains of more than 300 roe deer killed by
wolves were found in this year. In Naurzumsk preserve mass mortality
of roe deer was noticed after the occurrence of a strong snow crust
(16th March). This recurred in the snowy winter of 1948/1949. In the
winter of deep snow in 1947/48 wolves attacked even adult male wild
boars which they do not try to touch under ordinary conditions (Afanas’ev
et al., 1953). Frozen ground and protracted continuous snow cover* in
the steppes and deserts and the crusted snow in the forest zone show
similar effects. In all cases, the number of ungulates sharply decreases
and in subsequent years its significance in the foods of wolves de-
creases.

In some conditions, wolves feed on completely unlikely food. Thus,
during the mass outbreaks of locusts in the Kizlyarsk steppe in the
1920’s, the feces of wolves consisted entirely of the remains of these
insects (V.G. Heptner).

Home range. The way of life of wolves and their methods of
utilizing territory differ in different seasons and are not the same in
populations inhabiting different geographical zones and regions. Every-
where, two main seasons are distinguished, when wolves lead differing
mode of life. In the warm period of the year, in the time of parturition
and rearing the cubs, adult wolves live in pairs (families), while
pereyarki (young of the previous year) [yearlings] live singly or in

*In Russian, dzhvt—Sci. Ed.

162

222

small groups, usually not far from the parents. Both groups are fully
established or accomplish only short migrations mainly at the end of the
season.

By the cold time of year, the yearlings join the adults with this
year’s young. Thus a “pack” of wolves is formed, representing, in fact,
a family group consisting of the parents, juveniles (this year’s litter)
and yearlings which have not yet reached sexual maturity. It more
rarely happens that older animals, usually males 3—5-уеаг old, which
have been left alone for one or other reason, join such a “pack”.
Strange animals, born to other parents, are not admitted into the pack
and are regarded as enemies.

The average size of a pack is 1-2 adults, 3—6 juveniles, and 1-3
yearlings, i.e., 5-11 beasts. The largest pack known was composed of
16 beasts (2 adults, 10 juveniles and 4 yearlings). On the European
tundras, packs are usually composed of only the parents and young—
mostly 5—7 animals (maximum 10 wolves). Apparently, on the tundra,
yearlings do not always join their parents (Makridin, 1959). In
Arkhangel’sk district, the pack is usually composed of 7-8 and rarely
of 10—12 animals (Semenov, 1959). In wolf packs in eastern Siberia,
there are usually 7—10 animals (Khudyakov, 1937; Timofeev, 1949). In
Ryazansk district, in a total of 14 years of observations only once was
a pack of 11 encountered (1946), and 2 times of 10 wolves (1945 and
1947; Kozlov, 1955). In Tatariya, over several decades of observation,
no pack was found consisting of more than 12 wolves (V.A. Popov).
In Belovezhsk Forest, where wolves were numerous, a pack of 12
wolves was found in 1946. Apparently, in this region, packs of 15 and
16 wolves may be present, but more often, they consist of 8—10 animals
(Serzhanin, 1955). In Kazakhstan, 3—12 wolves occur in the pack, and
rarely—15. In the lower Ili river, packs of up to 10 wolves are met with
(Afansa’ev et al., 1953) (Table 15).

Table 15. Number of wolves in a pack in the lower reaches of Ili river

Number of wolves

Time of
observation

Average
size of pack

October-December 8 4 6 3 3 — 2 То. 2% 28 39
January 14 6 1 —- — — — — — Der 2h 1.4
February 2 3.30 о SS SS 8 Sil

*Figures in this column are clearly in error; they should be (from top down) 1,—,—.
—Sci. Ed.

223

„<< oO
"<. Копуаг\ ©,
-240 0.9090 Aa
“Uryadino®° ло о
) Ао
{=

>>>
|! о
о

Ie оо

< ante О
7 >>

„77ЗЫР>>
®

0°
xeon
nf

Lesozavod © iE
nt), ^%-
—о^

Tamarskoe la

So
10)
> | у
|
fo}
244“

“А
[®)

= о\— —
2 © Lupyazhi —" = = Dreserve™ vf
at 3 ‘Administration & 5

= Fe es 5
<® бтекроуот <<< Зе ВРиЯЛеУО °
°

° с 3
0° 8 Gorodkovichi °o
°.° ow

“Ay.
mM? о

1<<<4<<<<

‚Оегемепвкое == »>>;lvankhovos
>>>у? >>>555>>777 а я
>
аъ em = mae Kuchino
Rey ae aS ee a 95°
©

Fig. 55. Hunting areas of wolf pup litters in winter in Ryazan district in the
region of Oka preserve (from Kozlov, 1952).
1—location of litter, 2—hunting area of litters, 3—consistent passage of wolves,
4 ргезегуе boundaries, 5—migrations of wolves for prey in summer.

162

The average number of cubs in a pack in Aktyubinsk and North-
Kazakhstansk districts in December—January was 6.3 animals (of seven

packs; Afans’ev et al., 1953).

224

162 The pack is sometimes divided into groups. Thus, in the winter of
1944/45 a pack of 10 wolves, living in the region of Oka preserve, was
often divided into two groups, each of 5 wolves, hunting independently
at a distance of 8-10 km from one another. One group was led by the

163 mother and the other by the father (Kozlov, 1955). The winter pack
completely dissolves only with the deaths of both parents.

955

Коспетагуъ;

Ih"

|
|

Fig. 56. Distribution of wolf pup litters in the region of Oka preserve (Ryazan
district; after Kozlov, 1952).
1—place and year of litter, 2—preserve boundaries.

164

225

On the tundra, wolves feeding on prey from the traps of hunters,
carrion and bait at villages or edible flotsam by the sea, do not live in
packs, but singly or in pairs. Packs apparently cannot be sustained on
such food (Makridin, 1959). Old wolves incapable of reproduction live
also alone in winter (Kozlov, 1955). Sometimes, killing of weak or
diseased animals is observed in the wolf pack. Thus, wolves usually
tear apart and eat an animal which eats a strychnine bait and goes into
convulsions (Shvarts, Pavlinin and Danilov, 1951).

As arule, packs live separately and on meeting, they relate to one
another with hostility, although cases of serious clashes, fights, or
especially “battles” are unknown. Wolves which enter into the hunting
territory of a strange pack of wolves leave it at once upon their first
meeting with the owners of the territory, or upon discovering their
“scent marks” (see below). In rare cases, separate packs temporarily
unite and together attack a herd of wild and domestic ungulates; in this
case, the size of the united group may reach 20 and more animals. Such
a union is always transitory and of short duration, and usually happens
in places of crowding of ungulates—near large migrating herds, in the
places of mass calving, and so on. In the Mongolian Republic, such
joining of wolf packs was observed around large herds of dzeren

Fig. 57. Birch clear-cut in steppe—denning place of wolves. Krasnoyarsk
territory. May 1959. Photograph by V.V. Kozlov.

165

226

[Mongolian gazelle, Procapra gutterosa] (Bannikov, 1954). During
periods of pack life, wolves without growing young are very active and
in some places perform extensive wandering, following the migrating
wild ungulates or herded domestic livestock.

In the second half of winter and to its end, the mothers with their
by this time maturing yearlings separate from the pack and wander in
pairs or groups. The young wolves with the still not mature yearlings
in the lead, remain together, and in their absence, they disperse into
groups of 2—3 animals or wander singly.

The summer sedentary life of wolves begins, in the middle zone,
usually in April, with the arrival of pregnant females at den sites.
Whelping places (dens) are permanent; they are only changed in case
of extreme necessity, particularly alterations in the surrounding land-
scape unfavorable to wolves. The den is situated either at the center or
in one of the parts of the hunting territory of the family (see below).
Pregnant female wolves usually return to their old dens, while the
young that are pregnant for the first time, search for new lairs which
are, as a rule, not far from the place of their birth. At that particular
time the winter packs of wolves finally break up and the juveniles,
having now become yearlings, separate from the parents and live inde-
pendently, usually not far from the mother’s den.

Different wolf families have mutually exclusive hunting territories,
and the simultaneous living of two families in one territory 1$, as a rule,
impossible. Under certain conditions, at least in winter, this rule is,
however, not followed. Thus, in winter of 1920/1921, at the time of
great food scarcity (dzhut), a great number of horses wintered on
pasturage in glades in steppe areas and large meadows among the
clear-cuts of the Ara-Karagai forest (b. [former] Kustanaisk, Turgaisk
governance). The wolves, numbers of which were especially high in
that year, gathered thither from a large region and killed the horses.
The number and density of wolves in Ara-Karagai was so great that
there was no separation of hunting territories of the packs and perma-
nent packs were not really present (V.G. Heptner).

The territory of the pack and its boundaries are distinguished by
special characters—scent marks—places where wolves urinate. They
are usually situated near things—rocks, boulders, a single tree, the
skull or skeleton of a large animal—that are easily distinguishable from
afar. On passing by such a place, wolves must visit them and leave
their own marks, indicating that the spot is occupied. These “urine
points” are the best places for setting traps.

221,

Fig. 58. Small cliff in the forest-steppe under which was a den with 6 woif pups.
May 1958, town of Khakassiya, Krasnoyarsk territory. Photograph by
V.V. Kozlov.

The size of the hunting territory of the wolf family, and conse-
quently the distances between summer dens, depends on the availability
of food in the territory (stocks of summer food of wolves), cover, and
many other conditions. In the middle zone, dens are located usually no
closer than 6-12 km from each other (radius of the hunting territory is
3—6 km; Kozlov, 1955). In the Ukraine, summer hunting territories of
wolves are smaller. Dens here are situated near one another, sometimes
in neighboring ravines or in one woodlot (Korneev, 1950). Exception-
ally, two female wolves sometimes give birth to their young in a single
den. Thus, in the Sinyavsk region of Sumsk district in 1948, 8 wolf
pups and 2 old female wolves were found in one den. Among the cubs,
four were noticeably larger and obviously belonged to another litter
than the other four (Korneev, 1950).

In the taiga of eastern Siberia, the average radius of the hunting
territory is approximately equal to 10 km in summer (Khudyakov,
1937), while in steppe and especially in desert zones it is larger and
the wolves’ dens are located no closer than 15—20 km from one an-
other. Only a few exceptions are known. Thus, in the Vishnevsk region
of Karagandinsk district, three burrows of wolves were found at a
distance of only 500 m from each other. Two pups were found in one

166

228

burrow, 5 in another and in the third, 6 pups with eyes open were found
(Afanas’ev and others, 1953). In the middle zone, cases are known
(very rare) when two litters lived at a distance of only 2 km from each
other. These were 2 female wolves with their young, the males of which
(both) died in the spring (Sokolov, 1951). At the same time, there is
some indication that when neighboring dens are close, the female wolves
ate each other’s young (Danilov, 1945). In the mountains, dens are, as
a rule, located nearer to each other than in forest steppe and desert
zones.

At the time when the juvenile wolves are present in the den, the
parents usually do not hunt in the immediate neighborhood, but go out
to the farthest parts of their hunting region. Thus, in 1942 in the Altai,
in snow which had fallen on May 20th it could be traced that parents
went a distance of 12—25 km from the den for their prey, at a time when
wild ungulates and grazing livestock were living in their neighborhood

ee = 2 Е о

Fig. 59. Tracks of two wolves, passing a day lair, the place of their daily rest.
Khakassiya, Krasnoyarsk territory. 12 December 1956. Photograph
by V.V. Kozlov.

167

229

(Nasimovich, 1949; Afanas’ev et al., 1953). In the middle zone, the
radius of summer hunting chases does not exceed 5—10 km (usually 5—
6) (Sokolov, 1951). Thus, in Ryazansk district, in the period when the
young are being fed, the adult wolves hunt usually no closer than 5—
7 km from the den and do not touch livestock grazing within the limits
of this zone. In the summer of 1939 in the Charussk forestry [area], a
den of wolves with a litter was found 1.5 km from Chernaya Rechka
village; livestock from this settlement were pastured in the immediate
proximity of the place of the litter, but no losses caused by wolves were
noticed. At the same time, complaints about wolves were constantly
being lodged from those who were raising livestock at a distance of 7
km from the litter’s location. After destruction of the litter, there was
not even one case of attacks by wolves on livestock. Only yearlings
sometimes break the “rule” of not touching livestock near the den of
the wolf family (Kozlov, 1955).

In a convenient den, located in a favorable place, the litter lives the
whole time until autumn and the beginning of the nomadic period.
However, if the den is established by the adult wolves far from a water
source (2—5 km away), it is used only until the young are weaned from
the mother’s milk. Later, the litter resettles nearer water. The particu-
larly great need for water is manifested in the young at their transition
to a meat diet. Upon the drying up of a water source, wolves also lead
or transfer the young to a new den. During a period of drought in
Kazakhstan, wolf pup litters are concentrated near the few intact water
sources, devastating their neighborhood, while some litters perish due
to starvation and especially the deficiency or absence of water (Afanas’ev
et al., 1953).

A den-burrow built in open, non-overgrown ravines is also changed.
It is only used so long as the wolf pups do not come out of the burrow.
Later, the parents lead the litter to more hidden places (thickets of
shrubs and weeds, forest islands and sometimes to winter crop fields
where bread grain is grown)*. In the Oka floodplain, the adult wolves
frequently led the growing litters to places with an abundance of small
fruits—raspberries, strawberries, red billberry [Vaccinium vitis-ideae]
and others (Kozlov, 1955).

The she-wolf sometimes builds a temporary nest for whelping,
particularly when the places of the main nests are flooded by melt-
water, or when the she-wolf is frightened in some way before parturition

*Probably refers to winter wheat—Sci. Ed.

230

from the main nest. The duration of their stay in the temporary shelters
is determined by the local conditions, but before the young of the
current year become independent of their mothers, the litter usually
changes several temporary dens. In these cases, the she-wolf accompa-
nies the young to the main lair when they are already quite independent
(Sokolov, 1951).

If wolves in a den were disturbed, particularly if their litter was
touched, they usually abandon the den, carrying the surviving young to
a new place. With closing of the entrance to the den (burrow), the
parents usually dig it out at night and carry off the pups. At such times,
neither the female or male wolf defend the nest and do not attack its
invader, even if the nest is destroyed and the pups taken (Kozlov,
1955). While the adults are busy raising the litter, the yearlings remain
in the region of the litter, but they do not approach the den closely.
They live alone, or in groups of 2-3 animals and hunt independently.

Fig. 60. Resting place of wolf in forest-tundra. Photograph by V.P. Makridin.

168

231

At the start of raising the litter, the old wolves remain extremely
secretive; they usually do not howl in the region of the den until July.
In June in the south, and in July in the north, the demands of the young
for food noticeably increase, and the parents significantly expand their
hunting territory. Precisely at this time, the young begin to give voice,
apparently because above all they often feel hungry. After the young
develop, their elders begin to howl upon their return from the hunt, and
the juveniles answer the old ones. At the same time, cases of attacks
by wolves on livestock become more frequent in the vicinity of the den.
Thus, in 1938 in Kalininsk district, a horse was killed only 500 m from
a den, and the she-wolf for several days led the litter to the carcass to
feed. Near Bologi, a maturing litter of wolves assiduously tore up sheep
and calves, not farther than 2 km from their den (Sokolov, 1951).
Beginning with this period, together with the young (juveniles) may be
found the yearlings which join the family till the time of the next rut.
In July (in the south) and August-September (in the north) the maturing
young wolves attain half the size of the adults and begin to participate
in the hunt with their parents.

The size of the hunting territory continues to increase in autumn,
and the wolves gradually convert to a wandering life. Henceforth, the
old wolves stop bringing food to the den; on killing a large prey, the
parents lead the young to it, and after satiation, the family returns to
shelter by the morning (Khudyakov, 1937).

When the milk teeth of the young are replaced by permanent ones,
the wolf family begins to hunt together. At first, however, wandering
is restricted to the hunting territory. Roaming increases with the fall of
snow and the termination of livestock pasturing. In the middle zone, the
family rarely returns to the den site in the winter, establishing instead
diurnal lairs (Kozlov, 1955).

In forest regions of Arkhangel’sk district, from September wolves
already lead a nomadic way of life, making long movements and spend the
day where they are met by the dawn. At this time, each wolf family
occupies a definite hunting region, leaving it only in case of prolonged and
sharp hunger or continuous pursuit by humans. Tundra wolves are, at this
time, still more active. In eastern Siberia in winter, wolves wander widely,
following wild ungulates, mainly roe deer. Beginning in November, they
hunt ungulates almost exclusively; making chases or driving the roe deer
onto the ice, capturing them in their beds; and frequently bringing izyubr
[Manchurian wapiti] to “bay”. These predators do not remain for long in
one place; the hunting territory of the pack sometimes reaches a diameter
of 70 km (Khudyakov, 1937; Timofeev, 1949).

169

‘232

Fig. 61. Tracks of a pair of desert wolves, hunting a saiga. Central Betpak-
Dala, Kazakhstan. 7 January 1957. Photograph by A.A. Sludskii.

In the central districts, during snowy periods of the year, the hunt-
ing region of a wolf family is also expanded. In Ryazansk district, the
family hunting range of wolves had a diameter of 25 to 40 km. In forest
and forest-steppe districts of the European part of the USSR, wolves in
winter perform movements of about 15—30 km during the night and
when hungry, up to 50—60 km (Kozlov, 1955). In Voronezh and con-
tiguous districts, wolves leave the permanent summer habitats in
November, when the radius of their hunting activity does not exceed 5
km and begin to wander widely, dividing up into small groups, carrying
out invasions of livestock yards. At this time the radius of their hunting
range increases to 15 km and more (Mertts, 1953). In forest-steppe
regions, the greater part of the animals concentrate in forests, where in
winter the food is more abundant, and they usually occupy permanent
hunting ranges. With the coming of spring, they again wander back to
the fields (Mertts, 1953; Barabash-Nikiforov, 1957).

Pack formation in the snowy time of year makes it easier for
wolves to obtain food, especially the young (see below). In places with
deep porous snow cover, the pack moves in a line, always with an adult
wolf in front. In places with little or no snow, for example in the lower

233

Fig. 62. Tracks of a desert wolf following a saiga. Betpak-Dala desert,
Kazakhstan. 19 December 1956. Photograph by A.A. Sludskii.

Ili river, the pack frequently moves in a helter-skelter fashion (Afanas’ev
et al., 1953).

In searching for prey, wolves move around their hunting range
along a definite route (Fig. 55), using one and the same trails for
extended periods, restoring them after snowfalls; during this, the wolves
without fail find their old tracks, and follow them exactly, track by
track. Trails are laid out along the banks of rivers and shores of lakes,
in places of passage between plantations, ravines overgrown with shrubs,
etc. Widely used are roads and paths made by humans (Semenov,
1954).

Wolves on the tundra lead a pack way of life for the greater part
of the year, because attacks on [rein]deer herds and the pursuit of
stragglers from the herds are more successful when hunting in a group.

170

234

In the taiga zone also, particularly in the second half of winter, wolves
do not often find prey sufficient for feeding the whole pack, and thus,
cooperative hunting becomes temporarily unfavorable, and the wolves
hunt in pairs or singly, gathering in the pack only for diurnal rest.
Sometimes the pack is broken up and for a week or two the wolves live
in collections of not more than 2—3 animals (Semenov, 1954).

With the beginning of the rut, the mature wolves drive away from
themselves the young and yearlings that up till now have lived with
them. After heat which continues for S—10 days, and the end of rut, the
wolf family gathers together again for a short time for cooperative
hunting of large prey. In the north, approximately in April, such packs
gather on the main trails along which the [rein]deer herds move to their
summer pastures, and in following these herds, they move to the tundra
and to places of littering of the young.

Burrows and shelters. Different protected places serve as diurnal
lairs (resting places) for wolves. In cold, windy and damp weather, the
wolves prefer the more protected places, while in dry, calm and warm
weather, they willingly lie down in the open. In autumn—winter—spring
period, when the wolves lead a nomadic, or in general a more active
mode of life, they lie down wherever they are at daybreak. In summer
time, they are associated with dens which they must build for the young
(see above).

In constructing the den, the she-wolf usually makes use of natural
shelters—fissures in rocks, cliffs with overhangs by the banks of rivers
and slopes of ravines, holes thickly covered with plants, thickets of
saplings or bushes, “brushwood”, or fallen branches, weeds, etc. In
many cases the nest is constructed in the burrows of various animals—
foxes, badgers '(ш the forest zone), marmots (in steppes, montane
meadows), arctic foxes (in tundra), etc. Wolves usually widen the
adopted burrow and partly remake them. In rare cases, the she-wolf
digs a burrow, which is usually shallow and short, with 1-3 openings.
The burrows are built on hillsides with soft, mainly sandy and sandy
loam soil. In the majority of cases, the carnivore’s nests are situated in
dry sections.

The den, as a rule, is constructed not more than 500 m from a
water source (lake, river, creek, spring, deep hole with water, pond,
etc.); to it one-two trails lead, becoming especially noticeable when the
young begin to leave the burrow. The approach to the water source is
often concealed. Near the trails around the den, a large number of
resting places of wolves, places of play for the cubs, and food remains

171

235

are encountered. From the den there usually emanates a sharp odor. of
urine and of rotting food remains. The food remains attract such birds
as magpies, ravens and others to the nest. By their cries, it is possible
to locate with precision the nests of wolves with litters.

Convenient places for dens are few, and from year to year they are
occupied usually by one and the same pairs of wolves. If the male dies,
he is replaced by a yearling in the pair, and if it is female—by a young
she-wolf. Empty dens are therefore rare, and they are only left due to
the fundamental changes in the conditions of the surroundings (felling
of the forest, fire, marsh drainage, etc.). Thus, in the Charussk forestry
[area] of Ryazansk district, wolves established a den for four succes-
sive years (1936—1939) in one and the same woodlot, although every
autumn a drive was mounted against the litters appearing there, when
a portion of the young were killed, and in 1937 the mature male was
also killed (Kozlov, 1955).

The wolves more frequently select for dens, places rarely glanced
at by humans, but they construct them near domiciles, paved roads,
railways. In 1941 a wolf’s nest with young was found within 200 m of
the Moscow-Leningrad highway (Sokolov, 1951). In Zheltukhinsk re-
gion of Ryazansk district, a she-wolf built a den in part of a field
overgrown with weed only 1.5 km frorn the village. Another wolf den
was discovered 10 т from the rcad-bed on the Moscow-Ryazan rail-
way, under a pile of snow-screens (Kozlov, 1955). A she-wolf with a
litter lived in the middle of a peat-bog not far from a corduroy road
along which carts frequently passed (Manteifel’, 1949). In Ryazansk
district a den was found in a small but dense birch clear-cut not far
from a large village (regional center of Erakhtur) and in immediate
proximity to a large road with much automobile traffic (V.G. Heptner).
In the depth of large forest massifs, wolves do not usually make dens.
In Tatariya, all lairs were situated not further than 500 m from the
forest edge, usually along ravines where there is water (V.A. Popov).

In different landscapes, the situations and construction of dens are
somewhat different. Tundra wolves use shelters of three types: a) land
burrows, which they build themselves or adapted from ones of foxes,
and rarely, arctic foxes. The burrow consists of an entrance trench 1.5
m in length, an underground corridor 50—60 cm wide, and length 2 to
10 m, and a nest chamber 150 x 100 x 70 cm; the chamber is situated
at a depth of 1.5 to 3.5 m, and usually contains no nest bedding; b)
burrows under rocks of nearly the same construction, but usually shorter;
c) different natural shelters (caves, fissures in rocks, etc.). The wolf

236

Fig. 63. Tracks of a wolf and a she-wolf. Uzhursk region, Krasnoyarsk territory.
December 1957. Photograph by V.V. Kozlov.

den is usually situated on the southern and south-eastern slopes of river
banks or hills; they are better warmed by sun, and the snow thaws more
quickly there (Semenov, 1954).

On the Kaninsk tundra, wolves are mainly confined to hills in summer.
In Lena-Khatanga territory, burrows are usually no longer than 1.5 m and
not deeper than 1 m (A. Romanov, 1941). In Anadyrsk territory, wolves
give birth to pups mainly in land burrows (Sokol’nikov, 1927).

In the northern forest districts, wolves settle on dry ridges in the
middle of marshes, on islands of coniferous forest with abundant fallen
branches in coniferous saplings. They prefer burned-over mossy bogs,
especially in the difficult-to-penetrate margins overgrown with impass-

172

able small trees. Wolves settle in wet areas covered with alder woods
and on islands of dry, but dense, forests with spruce understory, which
are rarely visited by humans (Semenov, 1954).

Wolves living in the forests of Ural and Siberia, usually have dens
for their litters situated directly on the surface of the earth, under
bushes, overthrown trees, etc. With wolves of the open steppes and
forest-steppes, the denning place is more often built in a burrow, as a
rule in a foreign one—usually of fox, badger, or marmot (Khudyakov,
1937; Shvarts, Pavlinin and Danilov, 1951; Laptev, 1952). In the steppes,
burrows were also found distant from water sources—up to 12 km
(Borzinsk steppe). They were constructed in open, usually elevated,
places. In southeastern Siberia, a wolf den was found on a promontory
by some falls where only a few birches grew, but, it was dry there and
the she-wolf had a good field of vision, which protected it from surprise
attacks (Khudyakov, 1937).

In Belorussia, a litter of 6 pups was found in a fallen hollow alder
tree on a small hillock in a thicket of small alder trees (Serzhanin,
1955).

In the central chernozem regions, dens of wolves are situated in
young and old broad-leafed and mixed forest islands, in steppe ravines
and gorges overgrown with shrubs, and in reeds and willow thickets in
marshes, along the margins of bogs, and sometimes even in old straw
stacks and in the remains of corn-shocks. In the steppe section of
Voronezh district, the wolves raise their young mainly in “yarugas”—
ravines densely overgrown with small oaks. In forests, dens are
constructed under uprooted wind-fallen trees, sometimes in burrows of
foxes and badgers. In fields, the predators use old military trenches,
and in ravines—in holes and cavities.

In time past, there were repeated cases of pups being transferred
by she-wolves to tall cereal grains and construction here of dens in the
immediate proximity of villages (Mertts, 1953). In the Kantemirovsk
region of Kamensk district, in the autumn of 1953, was found a burrow
with two exits situated in a rye field where the wolves successfully
brought the young. Here such dens are not rare (Barabash-Nikiforov,
1957). In Chernigovsk district, a she-wolf built a den and brought the
pups to the hemp plot of a personal holding of a collective farmer 10 m
from his house. In the Kovertsovsk region of Volynsk district, a litter
of wolves lived in a rye field, 300 m from the cottage. The den was
constructed in a hole by the side of an uprooted stump. Such cases are
known in Sumsk, Zhitomirsk and other districts. In Donetsk district, a

238

wolf den was found 150—200 т from a very busy paved road (Komeev,
1950).

In the forest regions of the Ukraine, dens are frequently built on
islands among sphagnum and other bogs, in dense undergrowth of
saplings and in forest islands with dense shrubbery among fields
(Korneev, 1950). In forest-steppe regions, dens are situated in deep
forest ravines, in sparse forests along river valleys and in impassable
alder wood along tussock bogs (these places are called “volchatniks”
[belonging to wolves]).

In forest-steppe zone of Omsk district, the wolf selects for dens
ridges among marshes, ravines overgrown with bushes, thickets of sage-
brush in virgin land, and forest islands (Laptev, 1958). In the steppe
zone of the Ukraine, dens are more frequently present in deserted
gorges with thickets of spiny bushes and tall weeds in depressions
between sandy hillocks, overgrown by willow and other bushes, while
on the floodplain of the Dnepr and Dnestr—along thick stands of reeds
and other dense places (Korneev, 1950). In the forest-steppe and steppes
of Kazakhstan, den burrows are found in the forest, at the edge, and
in open places, usually among tall weeds or bushes at a distance less
than | km from a water source. In the forest preserve “Вогоуое”, a
wolf den was situated in a cave on a hill covered by a pine forest; this
den was thrice destroyed and near it, she-wolves were twice killed.

Dens dug by wolves in the steppe are usually 2—5 т long, rarely
more, with one or two exits, and the nest lies at a depth of 1-1.5 т
from the surface. Old, repeatedly used dens have very wide passages
through which humans can easily crawl, and several nest chambers;
entrances to the old chambers are frequently filled in when wolves
prepare a new one. Lairs in the open are established under cover of
bushes or tall weeds. In North Kazakhstan district an open lair was
found in the reeds of an almost dry lake; the nest was situated in a large
opening among meadowsweet [Filipendula], around which shallow water
was present. Within such a lake, a she-wolf brought a large heap of
cattail and reed and gave birth to 7 cubs on it.

In the desert zone of Kazakhstan, wolves give birth to cubs both
in burrows and in open-air dens. Burrows in the southern Pri-Balkhash’
were found in solonchaks [salt flats]. They were 7 and 8 m long and
1—1.5 т deep. In sandy mounds covered with saxauls, a den was
excavated at the top of a mound under the roots of saxaul and consisted
of a simple niche, in which 6 cubs were lying. One burrow was situated
in a dense growth of saxauls in the Ш river valley. Here open-air lairs

173

239

were also found in reed growths and under willow bushes (Afanas’ev
et al., 1953). In Kazakhstan, of 24 wolf dens, 14 were situated in
burrows dug by she-wolves, one was in a badger burrow, another one
in a cave, another in a niche and 7 were in the open (in tall weeds, rose
willows or reeds).

In Turkmeniya, wolves inhabit the plains, old stream channels in
cliffs of the chink along the Uzboi, in ravines of the Kopet-Dag [moun-
tains], near creeks and springs, chiefly in the shady places, under the
overhanging rocks, etc. Sometimes, dens are situated in dense growths
of tamarisks (Tamarix) or other bushes, and often in tugai (riparian
vegetation) (V.G. Heptner). In Armenia, these predators usually use
narrow caves, fissures and ravines for dens (Dal’, 1959).

In the montane steppes of Kirgiziya, on high-alpine watersheds and
in the foothills, wolves usually construct the den in a burrow dug in a
ravine, under a bush, or in another such protected place. On the slopes
of mountains, they use fissures in the rocks, overhanging shelves of
stone, marmot burrows, etc. (Kuznetsov, 1948).

Fig. 64. Tracks of a desert wolf in mud near a waterhole in the Gyaz’-Gyadyk
mountains. Badkhyz preserve, southern Turkmeniya. February 1965. Photograph
by V.G. Heptner.

174

240

Daily activity and behavior. Wolves are nocturnal predators. In
the majority of places they lead a nocturnal mode of life in summer as
well as in winter.

In winter, wolf litters begin hunting in the early evening twilight; they
hunt the entire night, sometimes traveling tens of kilometers, but neverthe-
less often they lie down to their diurnal rest hungry. In sparsely settled
regions, hunting during the day is also observed, usually for wild ungulates
(Korneev, 1950). In those places where wolves are active only at night,
individual wanderers also hunt during the day. Thus, in January 1951, in
the region of the Oka preserve, one yearling she-wolf picked out among all
prey particularly those that were active by day (V.G. Heptner). Attacks of
wolves on livestock were common, and especially on single horses during
the daytime, often at the village itself, in the period of their very high
numbers at the beginning of the 1920’s in former Kustanaisk district of
Turgaisk governance (V.G. Heptner). With the arrival of winter, when
lying down rather than a bed the wolf turns around until there is no longer
any packing of the snow, in order to form a good pit-lair. While sleeping,
wolves space themselves in a semicircle, covering their “scenter” (end of

Fig. 65. A wolf calmly walking in deep snow. Krasnoyarsk territory. January
1958. Photograph by V.V. Kozlov.

174

241

the nose) with their tail. Sleep is very light, the ears are constantly either
pricked up or drooping. Wolves sleep especially lightly in times of frost.

In eastern Siberia, in summer, the wolves mainly stick to a noctur-
nal mode of life, although whelps are frequently active by day also. In
autumn, after the transition of the whole litter to a nomadic life, they
hunt mainly at night.

In southern Turkmeniya (Badkhyz), wolves lie by day in tamarisk-
like or other shrubby thickets, and sometimes almost at the outskirts of
settled places, always relatively close to water sources, but in sandy
areas, at a distance of 15 km from wells (Heptner, 1956). In settled
regions of Kirgiziya where wolves are frequently disturbed, they mainly
lead a nocturnal mode of life; but in remote places, one may see them
hunting during the day (Kuznetsov, 1948). Mongolian wolves hunt at
any time of the day or night (Bannikov, 1954).

Hungry wolves become aggressive. A huge number of cases are
known of severe attacks of wolves on livestock. Thus, on | February
1953, two wolves attacked the calf-pen of a collective farm in the
Dryazginsk region of Lipetsk district. On hearing the noise, the people
came running; however, their appearance did not cause the wolves to
run away. A struggle began which ended only after one wolf was struck
with pitchforks. In autumn, wolf litters commit audacious onslaughts on
herds of sheep and flocks of geese, sometimes seizing their prey almost
from the hands of the shepherds. Under ordinary conditions, wolves
fear humans and are very cautious. They can well differentiate between
dangerous hunter and unarmed herdsmen, in front of the eyes of which
they frequently attack the sheep.

Pursued wolves sometimes seek shelter in villages. In 1950, a wolf,
having been shot from an airplane near the village of Vyselka
Rozhdestvensko—the Khavsk region of Voronezh district, fled to a
village and took shelter in the hayloft of a house near which a calf was
lying. The wolf was discovered only after some time (Barabash-
Nikiforov, 1957). In Kirgiziya, the wolves in winter approach villages
and frontier posts even during the day (Kuznetsov, 1948).

Wolves hunt in various ways: from concealment, pursuit of prey,
and organized as a drive in groups. In summer they hunt mainly singly,
the wolves frequently ambushing their prey, rarely spotting and pursu-
ing prey. When attacking a herd, even individually, wolves try to frighten
the animals, creating panic and dispersing the herd or separating one
or several animals from it.

175

242

Upon attacking a herd, wolves are able to distract the attention of
the shepherd and often conduct a very well-organized combined hunt of
ungulates and hares, in which the functions of each member of the pack
are not only different, but also coordinated. Breaking off from one
another, they envelop the pursued animal from different sides and at
last surround it. Frequently, some of the wolves pursue an animal and
another group tries to intercept or set up an ambush in its path (Timofeev,
1949). Once on the Malyi Cheremchan river in Tatariya, there was
observed about 9 in the morning a unique hunt by wolves for geese: a
wolf drove the swimming geese to a bridge, at which a hidden she-wolf
intercepted them (V.A. Popov).

Along black [snowless] paths or in light snow, wolves can pursue
prey at a speed of 55—60 km per hour; with this they quickly catch even
the brown hare [Lepus europaeus] which runs at a speed of 40-45 km
per hour. Wolves obtain roe deer by driving them in persistent pursuit.
They destroy many marals by chasing them in winter into [soft] sedi-
ment surrounding a rock, the predators patiently wait until the legs of
the maral stiffen and it becomes unable to remain by the stones. Four—
five wolves do not fear to attack an elk on frozen snow-crust, but with

little snow, the adult elk is for them a dangerous enemy (see above).

The pack can easily cope with young pigs and sows, but they usually
fear to attack boar (Timofeev, 1949). Wolves pursue and frequently kill
foxes but they rarely eat them. Hence, where there are many wolves,
foxes usually become few. In the European part of the [Soviet] Union
wolves in winter very successfully lure dogs from villages and catch
them by feigning a cowardly flight (Korneev, 1950). Wolves at night,
and sometimes during the day, enter villages and settlements. Thus, in
August 1954, wolves carried away a sheep which spent the night in the
street from the outskirts of the village of Kopanovo (Ryazansk district),
and injured another (V.G. Heptner). Wolves follow the barking of the
hound dogs for many kilometers and often catch them during the hunt-
ing season. In the agricultural central-chernozem [black earth] regions,
wolves have adapted to “mousing” behind the tractor ploughs, remain-
ing some distance from it and catching the voles flushed out by the
plough (Mertts, 1953).

In searching for prey, the wolves depend mainly on their sight and
hearing; scent is relatively weakly developed in wolves, and therefore,
they rarely discover hidden hares or birds. But, when they come across
a fresh track, they can easily follow it without becoming confused
(Timofeev, 1949). Hearing of wolves is very sharp; the organ of hear-

176

175 Fig. 66. Diagram of the tracks of a wolf

243

ing is constantly exerted, the wolf hears and constantly registers the fall
of autumn leaves from the trees. It nearly never happens that a sleeping
wolf can be approached, while it is not very difficult to approach a
sleeping fox. Only a replete sleeping wolf can sometimes be approached
to within 10—30 steps.

The sense of smell of the wolf is weaker than that of many hunting
dogs. The wolf can scent carrion on the wind for not farther than 2—
3 km. Because of the comparative weakness of their sense of smell,
wolves can be taken in traps relatively easily. The sight of the wolf is
not so complete as that of dogs. In darkness, the wolf sees better than
the other species of the dog family because, through their way of life,
they are by preference nocturnal animals (Korneev, 1950). Even at

2 an
sys 2% 4 374

pack walking, as usual, one behind the other; 6
when turning they separated but again went

one behind the other. The tracks during the Я
turn represent the only reliable possibility to 6
establish the number of wolves in the pack— у
4 т the present case (from Kozlov, 1955).

%ь:

244

Fig. 67. А wolf on- guard. Khakassiya, Krasnoyarsk territory. Winter, 1957.
Photograph by V.V. Kozlov.

night the wolf can easily detect the flaws in disguised traps. At the
same time, wolves are far-sighted and they can distinguish between the
armed hunter and the simple passerby at great distances. Upon concen-
trating their attention on any item or occurrence, the wolf considerably
blunts its perception of others. Numerous occasions are known when,
fleeing from beaters, wolves came very near to shooters, even those not
hidden. A wolf may approach with the wind, a person standing motion-
less especially when dressed in camouflaged clothing up to 5—10 т, not
smelling an odor or noticing a motion (Kozlov, 1955).

On the whole, the level of nervous activity of wolves, and their
ability to consider situations is quite sufficiently high. The behavior of
wolves is more complex and “wiser” than is assumed by the majority
of zoologists and hunters. When going to their dens, adult wolves
skillfully select hidden approaches. At that time, they make numerous
“dodges” and “double-backs” using bushes, wind-fallen trees, stumps
and many others. Their ways of hunting, as shown above, are highly
variable and they are usually governed by the complex and highly
coordinated behavior of the members of the litter (pack). The complex
nervous activity of wolves with the highly developed and multiple
conditional reflexes (temporal connections) allows them to adapt them-

177

245

selves quickly to new situations. This “flexibility” in the behavior of
wolves, for example, their ability rapidly to distinguish poisoned baits
and avoid them, makes their control difficult.

Seasonal migration and transgressions. Wolves in the majority of
regions are settled. They abandon their accustomed places quite unwill-
ingly, usually only in cases of severe shortage of food in their hunting
territory. The migration of wolves is, in the majority of cases, con-
nected with nutrition and represents nomadic movements for food.

Significant seasonal movements of wolves have a place only in
tundra, steppes, deserts and montane regions with nomadic (shifting)
livestock breeding, where the herds of reindeer, horses and cattle are
regularly driven from the summer pastures to winter and vice versa.
Examples of such nomadism are performed by herds of reindeer in
tundra, saigas, dzerens and goitered gazelles in steppes and deserts, roe
deer in the Urals and in Siberia, and by deer in the mountains. The
times and paths of migration of the ungulates and the wolves coincide.
In mountains, during the fall of deep snow, wolves, following the
ungulates, move to less snowy places, usually descending to the foot-
hills.

In the tundra wolves lead a nomadic way of life for three-fourth of
the year, not adhering to stable hunting regions. Twice yearly they
accomplish large meridional migrations connected with the driving of
the [domestic] deer herds. When the deer herders shift southward, the
wolves go together with the deer herds to the forest border. They do not,
as a rule, penetrate deeply into the taiga, although in the European north
in particular, the main mass of the deer herds of the state farms and
collective farms winter there. Wolves prefer to remain in the forest-
tundra and mossy bogs with less deep and more firm snow cover. Here
in the forest-tundra, there is much natural food, since willow ptarmigan
migrate thither, white hares are very frequent, and until recently, elk
were encountered along the river valleys where trees are located
(Semenov, 1954). In the Nenetsk Nats. Region, besides seasonal migra-
tions, the movements of wolf packs from the Bol’shezemel’sk to
Malozemel’sk tundra in connection with herd migrations were noted,
but return movements were not (Makridin, 1959). Seasonal migrations
of tundra wolves in the European north reach 200—300 km (Semenov,
1954).

A small portion of the wolves remains on the tundra in winter.
They move out to the sea coast, where remain only small reindeer herds
of hunters engaged in the arctic fox harvest and near fishery stations,

178

246

feeding on fish and other animal wastes. On the seashore wolves find
the bodies of stranded marine animals. Tundra wolves are very mobile
and move for great distances. A trailed pair of wolves went 70 km in
one night (Semenov, 1954); from an airplane it was possible to follow
the path of a pack of wolves that went more than 150 km in 20 hours
(Semenov, 1954). Regular migrations to the Novosibirsk islands, fol-
lowing reindeer across the sea ice are accomplished by wolves in
northern Yakutiya. In the forest zone, wolves are less mobile. They
follow roe deer during their migration in the taiga of eastern Siberia
(Khudyakov, 1937). During food abundance, the predators remain for
5—8 days in comparatively small territories; if they do not obtain food,
the wolves go “on foot”, moving 25—50 km in a 24-hr period.

In the montane forest landscapes of the southern Urals, wolves
migrate with the ungulates in winter from places of deep snow. In
Bashkir preserve wolves, together with roe deer and marals, remain
in the winter in the montane massif of South Krak and stay in small
numbers in the Uraltau (Kirikov, 1952). In the Altai, they leave the
mountains following the roe deer into the lower valleys. In the Tien
Shan, the predators ascend in summer into the mountains (to high
watersheds) following the herds of wild ungulates and domestic ani-
mals and they again descend lower when deep snow falls in the
mountains.

Generally, settled populations of wolves live in Caucasian,
Borzhomsk and Zakatalsk preserves, as well as in the reed beds of the
Caspian coast; however, they also perform vertical and horizontal mi-
grations following the wild ungulates. In the remaining territory of the
Caucasus, especially in its eastern part, wolves wander after the herds
of domestic livestock, ascending the mountains in summer and in
winter—to the lowlands (Vereshchagin, 1959).

Wolves accomplish regular migrations in the steppe regions, where
transhumant animal breeding exists or where large herds of wild ungu-
lates survive. Following herds of dzeren [Mongolian gazelle] on the
[USSR] boundary in the Trans-Baikal steppe, wolves frequently pen-
etrate from the Mongolian Republic. In the deserts of central Kazakhstan,
these predators accompany the herds of saigas and goitered gazelles
going, in September—October, to wintering grounds in the southern part
of Betpak-Dala. From the Severo-Kazakhstansk district, the majority of
wolves, in December—April, leave for more southerly regions (Afanas’ev
et al., 1953). In the Kustanaisk steppes, in the periods when wolves
were abundant in 1920—1921, in autumn they moved about, following

247

the herds of horses and going by day behind the horse drov-
ers at a distance of only few hundreds of meters or even nearer (V.G.
Heptner).

In deserts and in part in the steppes, the cause of wolf migrations
may be steppe fires, drought and lack of water or other reasons leading
to massive translocation of livestock and the migration of wild ungu-
lates. In connection with fires in the Badkhyz desert and consequent
lack of water, goitered gazelles appeared en masse along the Kushka
river in June 1942. Following them thither, the wolves appeared. This
massive appearance of gazelles, and of the wolves following them was
noted in 1927 along the Murgab river. In other desert-steppe regions
of such a sort, migration of desert and steppe ungulates to waterholes,
in the middle of summer chiefly in the river valleys, are always accom-
panied by corresponding migrations of wolves (Heptner, 1956).

Besides migrations for food, usually taking the character of sea-
sonal migrations, wolves sometimes perform long, time-consuming
massive migrations not having a regular character. They are either
connected with unfavorable changes in their principal habitats, or rep-
resent the growing cubs establishing themselves.

In some years, it was noticed that tundra wolves migrate in a
horizontal direction. Migration of wolves was observed from the Urals,
to the Yamal, into the Bol’shezemel’sk tundra; the wolf packs crossed
the Malozemel’sk and Timansk tundras and reached the Kanin isthmus.
Such migrations of wolves usually coincide with massive migrations of
arctic foxes in that direction. A return movement of wolves (from west
to east) was not observed (Semenov, 1954).

In Kazakhstan in individual years wolves also perform long and
irregular movements. Thus, in January 1941 in Dzhambulsk district
many “Siberian” wolves appeared, which were well distinguished from
local ones by their large size and gray color. In autumn 1942, wolves
moved en masse from the northern Caucasus, from Rostovsk and
Volgogradsk districts into Zapadno-Kazakhstansk, Gur’evsk and even
Aktyubinsk districts of Kazakhstan. The large European wolves well
differentiated from the small, reddish local ones (Afanas’ev et al., 1953).

Settlement of wolves is easily registered when they enter into places
which are unsatisfactory as habitat. Thus, during the summer, wolves
penetrate into the taiga of western Siberia from districts comparatively
far to the north and south, that are continuously inhabited, for tens or
even hundreds of kilometers (Aleksandrovsk region of Tomsk district,
upper reaches of the Taz river, to Surgutsk region; Leptev, 1958).

eh)

248

However, when the deep snow falls, they abandon these places or die
there. Wolves which in winter inhabit the Berezovskii region of
Tyumensk district along the Syn’ and Lyapin rivers, migrate in summer
to the Urals (Flerov, 1933).

Under difficult conditions lengthy migrations are known. Thus,
from 1933 to 1945, wolves four times penetrated over ice to Barsa-
Kel’mes Island in the Aral Sea which lies 80 km from the shore
(Afanas’ev et al., 1953). In 1938, three wolves walked on ice about
200 km and reached Wrangel Island.

Reproduction. Wolves are monogamous; the united couple usually
stays together for life, if one of the wolves does not die. Upon the death
of the male or female, as shown above, pairs are quite quickly re-
established. Since in wolf populations, males often predominate (Teplov,
1948), unpaired females are a rare occurrence, while unpaired males
are quite common; this is related to the later maturity of males. Among
wolves caught in traps and from airplanes in the European tundras,
71% were males in 1951-1953 (31 wolves studied); in 1954, 67%
males (30 wolves) and in 1955, 50% (72 wolves); the average of all.
years (133 wolves) was 59% (Makridin, 1959).

Wolves mature at about 2 years; practically all she-wolves become
sexually mature at the end of the second to beginning of the third year
of life; males in part begin to participate in reproduction in the third,
and in part the fourth year of life. Estrus in she-wolves and rut begin
usually at the beginning to middle of the second half of the winter. Old,
multiparous she-wolves enter estrus first, approximately 2—3 weeks
earlier than heat in young, just maturing she-wolves. Before the begin-
ning of rut, as was shown, the wolf pack dissolves. Adults disperse first,
then yearlings, and finally juvenile wolves. Later on, they re-unite, but
they do not approach the adults and lie separately. Driving away the
young, the adult male becomes unusually attentive to the she-wolf, does
not leave her for a minute and opens up a path for her in the snow.

Heat in older animals usually proceeds calmly, but in certain (rare)
cases it becomes complicated dué to the presence of 1-2 unpaired
males. Heat lasts approximately 2 weeks. At the beginning of heat and
rut in the young wolves, they drive away the juveniles still living with
them at that time. Particularly at this time, during the formation of new
pairs, the so-called “wolf wedding” is manifested, when a group of
males gathers around one female. These “weddings” are accompanied
by violent struggles and hard fights by young males for the female. The
fight sometimes ends with death and the dead wolf may even be

180

249

devoured. Such a group does not remain in one place for long, but
wanders widely spending the day in remote places—wilderness, bogs,
fir-groves, often wandering the byways, usually at night but sometimes
also by day time. The she-wolf bonds with the strongest wolf of this
pack and thus a new pair is formed, lasting until the death of one or
both “spouses.”

The periods of estrus and rut vary in different regions. In the south,
copulation takes place 1-1.5 months earlier than in the north; therefore,
the period extends from December to’ March. In the Karelian ASSR,
estrus is initiated at the end of February and in March (Marvin, 1959).
In Arkhangel’sk dist[rict], rut begins at the beginning of March in the
northern regions, and at the end of February in the southern, while in
regions adjacent to Vologodsk district, it begins approximately one
week earlier (Semenov, 1954). In the forest-tundra and tundra of Nenetsk
Nats. Region, the first female in which ovulation had taken place was
caught on the 9th of March. In the period 9 March—14 April, only
yearling females were still not ready for the rut (Makridin, 1959). In
Tatariya, rut usually occurs in the middle of February. In the middle
Urals, wolves copulate in February (Pavlinin, Shvarts, Danilov, 1951).
In forested montane landscapes of the southern Urals, estrus in wolves
proceeds from the end of February—beginning of March (Kirikov, 1952).
In the former Tarsk region, estrus in wolves proceeds from the second
half of January—first half of February, and whelping in April (Shukhov,
1928; Kuklin, 1938). In Irkutsk district, estrus begins in wolves at the
end of January (Timefeev, 1949).

In Belorussia, rut is initiated at the end of January; it ends approxi-
mately at the middle of March (Serzhanin, 1955). In Voronezh district,
rut in adults begins at the first, or more often the second third of
January. At this time, the characteristic howl of wolves can be heard
around dawn, but two weeks later, the breeding pack of one female and
2—3 (here rarely more) male wolves can be found during day. Rut ends
at the beginning of February, but sometimes extends to the first half of
March. After that, the formed pairs occupy territories, and, in April, the
she-wolves give birth (Mertts, 1953; Barabash-Nikiforov, 1957). In
Alma-Atinsk district estrus in she-wolves proceeds from December—
January. In the lower Ili river, a she-wolf in a state of estrus was
caught on 27 January 1949. In northern Kazakhstan, estrus occurs in
February—first half of March.

The duration of pregnancy in the she-wolf is 62—75 days.

In Nenetsk Nats. Region, the largest number of embryos occurring

250

in pregnant she-wolves was 9; the average number of embryos was 6.5
per female. From one to nine pups have been found (6.7 per female on
the average) (Makridin, 1959). The number of whelps in a wolf litter
in Yamalo-Nenetsk Nats. Region is more often 3—4, rarely 5. The usual
number of a pack (litter) is 7-9 wolves, including 2-3 yearlings

Fig. 68. Uterus of she-wolf with 9 embryos and uterus of a young nulliparous
she-wolf, [both] caught in March. Photograph by V.P. Makridin.

—

180

(Rakhmanin, 1959). In the middle Urals, there are up to 12 young (up
to 14 fetuses were found), more frequently 5—8 (Pavlinin, Shvarts,
Danilov, 1951). Young she-wolves produce smaller litters, usually 3—
5 young; adults are more fecund (6—8 and up to 14 young). Old females
in Belorussia produce 8 young, rarely more, and young—3-—54, giving
an average litter size of 4—6 pups (see Tables 16 and 17).

The average size of litters varies in different years.

Before whelping, the she-wolf usually returns to the old den in
which it has raised the past litter. A young one, whelping for the first
time, searches for a new den, usually not far from its birth place.

Parturition of pups takes place in the warm months, when the
stocks increase and food becomes varied. In Karelia, 3—7 (rarely more)
blind pups are born in April-May or the beginning of June (Marvin,
1959). In Arkhangel’sk district, forest wolves whelp at the end of April
and the first third of May. They give birth to 3-12 pups (more often
5—8). In the tundra, pups are born around the second half of May—
beginning of June; the young are from 3 to 8, 6 on the average (Semenov,
1954). In the southern Urals, pups appear at the beginning of May. On
21 April 1946 a killed she-wolf had embryos of about 310 gm each
(Kirikov, 1952). The earliest litters are found in Belorussia in the
second half of March, more often in April and May. In Brestsk district
in 1948, pups still blind were found on the 10th, 22nd and 27th of May.

In the central chernozem districts, there are usually 3—4 pups in the
litter. In 1950 litters of 6 and 8 pups were captured, and in 1956 a litter
of 10 pups. The low average [3—4] litter size in these districts can be
explained by the fact that the majority of females die when they are still
young, far from reaching the age for which a high litter size is char-
acteristic (Mertts, 1953).

In the Ukraine the pups are born at the middle of March—-beginning
of April. The earliest litter was found on 3 January 1949, of 5 one-
week-old pups (Kiev district). In Sumsk district, two pups approximately
3 weeks old were found on 2 June 1947. In the first third of June, a

Table 16. Frequency of occurrence of wolf litters of various sizes in
Belorussia (Serzhanin, 1955)

Number of 1 24S бло. п 1 otal
pups in litter
Number of 5 Suede Ол 6 7 — 1 — 1 65%

litters
SSSR BETES SUSE SENS SEE ees AVR SE re AE ote REED,

*Should total 64—Sci. Ed.

182

Table 17. Average number of pups in a den (Serzhanin, 1955)

Years 1947 1948 1949 1950* 1951 1952
Number of dens 81 186 150 168 180 173
Average number of 4.5 4 4.5 92 4.9 4.2

wolf cubs in den

*In Russian original, 1959—Sci. Ed.

Table 18. Frequency of occurrence of wolf litters with different numbers of
young in Kazakhstan (Afanas’ev et al., 1953)

Number of pups
in the litter Dre OA SAINT) eo О 1213 то

Total number

of cases 7109 ее об
Including:
In southern
districts 2 DB Вы eal

In northern and
central districts 2 By 55) 8 6 9 5 6 2 5 2 1 54

litter was found in Chernigov district, and the latest litter was found in
Volynsk district in July 1949 (Korneev, 1950). In Zakarpatsk district
with its mountainous climate, in the majority of cases birth of the young
usually takes place at the end of May. In individual cases, in the
Ukraine, litters attain 10—11 young.

In the north and the central districts of Kazakhstan, small pups
were found on 16 April (2 litters), from 2 to 30 May (17 litters) and
in June (1 litter). From 2 to 13 pups occur in litters, the average litter
size in the southern districts of Kazakhstan being 6.3 pups and in
northern and central districts, 7.0 cubs (Table 18). A litter of 13 pups
was found once, the mother and pups being in extreme emaciation
(Afanas’ev et al., 1953).

In southwestern Turkmeniya, the she-wolf produces young at the
end of March—beginning of April; on 12 April 1942, blind pups 5—7
days old were caught. In the litter 3—4 pups usually occur (Dement’ev,
1959). In the Caucasus, the she-wolf gives birth to 5—6 pups on the
average. Time of birth of pups extends from March to May (Satunin,
1945). In Azerbaidzhan, litters appear in April-May (from 2 to 8
young) in secluded places—remote ravines and in dry islets in marshes
(Vereshchagin, 1951). In Armenia, wolves copulate in January, giving
birth to cubs in March; the litter is 3—8 young (Dal’, 1954).

253

Fig. 69. Wolf pups 10-11 days old, taken away from the den. Khakassiya,
Krasnoyarsk territory. 18 May 1956. Photograph by V.V. Kozlov.

Maternal sense of wolves is very strongly expressed. A case is
known when a she-wolf followed a hunter who had captured its cubs
into a village and at night walked near his house for a long time.

Out of 5—7 pups (average litter size) born, only 1—3 animals reach
sexual maturity (juvenile mortality is 60-80%). Wolves usually live
not more than 15 years. A wolf which lived for 16 years in captivity
was quite decrepit (Afanas’ev et al., 1953).

Growth, development, and molt. Wolf pups are born blind, with
closed ear openings, and covered with short soft grayish-brown fur.
The weight of new-borns is 300—500 gm. They begin to see on the 9th—
12th day; at the end of the second—beginning of the 4th week, milk
canines erupt. In captivity, pups which begin to see on the 9th—14th day
grow strong and healthy, while those in which vision develops later
remain usually sickly and ricketic. 3-week-old pups crawl out of the
den, and at 1.5 months old, they quickly flee from danger and take
cover. Adult wolves leave their 3-week-old cubs alone when setting
out on a hunt.

In the first weeks, the mother does not leave the litter and during this
time the father-wolf feeds her, bringing prey to the den, or regurgitating

183

254

the semi-digested meat. The she-wolf nourishes the young with milk
until they are 1.5 months old. When the pups are 3—4 weeks old, they
begin to eat the regurgitate offered by the father, and the mother only
eats the remains. The necessity of regurgitative feeding (semi-di-
gested meat) is, apparently, related to deficiencies in digestive juices
(peptidases) in the young whelps. Artificially fed pups without such
regurgitate are accompanied by poor growth, delayed development
and rickets. Pups 3—4 months of age already find such regurgitate
unnecessary. In summer, nursing she-wolves become very thin. Milk
gradually becomes, for the young, only a supplementary food, and
they feed mainly on small animals which the father and mother bring
them.

In the first four months, the tempo of growth of the pups is high.
During this time, their weight increases nearly 30 times (from 350—
450 gm to 14-15 kg; Novikov, 1956). In the southern Urals, the
weight of a juvenile male in August 1941 was 11 kg, of a juvenile
female—8 kg, in September—13 kg. In the next 3—4 months, the
weight of wolves doubles and in December, they reach 28-30 kg
(Kirikov, 1952; Gavrin and Donaurov, 1954). In Ryazansk district,
young at 6 months attain a weight of 16—17 kg.

Later, when the pups become strong, the mother and father bring
living, but injured, prey to them in the den, teaching the whelps how
to catch and kill it. At the middle of summer, the adults begin to lead
the pups to killed prey. The actual training of the young begins at the
second half of the summer. From August, they attempt to catch rodents
and other small animals by themselves, and starting from September
they share with the adults in hunting ungulates.

The molt of wolves takes place twice per year; in spring from the
end of March or middle of April to July. In autumn there is no full
replacement of the pelage, only the underfur growing and developing
(from the beginning of September to the end of October, and sometimes
to the end of November).

Enemies, diseases, parasites, mortality, competitors. This is little
studied. The ticks Ixodes ricinus, Dermacentor pictus, Sarcoptes scabiei
[the mange mite] and others parasitize wolves in great numbers. In
distinction from foxes, only a little full-blown mange is ever observed
on wolves; it more often strikes young wolves. Among other ectopara-
sites may be noted: the biting louses Trichodectes canis, the sucking
louse Linognathus (sp.), and the fleas Pulex irritans and
Ctenocephalides canis (Korneev, 1950). It is completely obvious that
this list is very incomplete.

184

255

Among endoparasites are established the nematodes Тохосага
leonina, T. canis, the cestodes Taenia pisiformis, T. hydatigena, Echi-
nococcus granulosus, Mesocestoidea lineatus, Dioctophyme renale
(through fish) and the adult phase of the causative agent of sheep
gid[diness]—Multiceps multiceps. Infections with helminths are not
great. In the central chernozem districts, of 20 wolves examined, nema-
todes were found in the stomachs of seven (4—11 specimens) in the
small intestine of one—many fine cestodes, and in the trachea—5
nematodes (Barabash-Nikiforov and Pavlovskii, 1947).

Wolves frequently suffer from rabies and they act as a source of
this infection for people. The overwhelming majority of “attacks” of
wolves on humans stem from rabid wolves which, because of the dis-
ease, lose the caution and fear of man of normal wolves. Although
70-88% of the total cases of rabies among humans is due to bites by
dogs, cats—6—8%, and wolves—tess than 2—3% (Rudnev, 1950), wolves
play the role of principal host (reservoir) of the rabies virus in nature
(Zhdanov et al., 1955).

The only serious enemy and competitor of wolf is, apparently, the
tiger. In the Ussuri territory, wolves were absent in some places from
time immemorial and they appeared only after tigers were extermi-
nated. It was also noticed that wolves disappear from those places in
which tigers appear.

The mortality of wolves is great. In the course of the first year of
life, as shown, it reaches 60-80%, and subsequently decreases to
approximately 40-50% per year. Because of this, young of the cur-
rent year usually compose about one third of the population (Makridin,
1959).

Population dynamics. The number of wolves, though relatively
stable, changes under the influence of different conditions of existence.
It is necessary to accord the first place for significance their pursuit by
humans, which increases with increase in density of settlement and the
development of agriculture; it becomes weakened when agricultural
activity is disturbed for one reason or another. Therefore, a rise in the
number of wolves was observed after each war, famine or economic
dislocation.

At the beginning of the nineteenth century, wolves were numerous
in the majority of the developed regions of the country, including the
entire Ukraine both in Poles’e, and in the steppes of the Black and
Azov seas. The abundance of predators in the forested parts of the
Ukraine was related to the abundance of wild mammals (deer, roe deer,

256

wild boar, beaver, hares) and birds. The number of wolves particularly
increased after the French invasion in 1812 when the numerous bodies
along the route of the French retreat served as food for wolves. At that
time, there was even noted a migration of predators to Ukrainian Poles’e
from neighboring districts of Belorussia (Korneev, 1950). At the middle
of the nineteenth century, although the populated areas in the Ukraine
increased significantly, wolves remained numerous even in the vicinity
of such large cities as Odessa. In only 4 days in the autumn of 1844
during a mass drive in [the former] Khersonsk, Ekaterinoslavsk,
Tavrichesk and Bessarabsk governances, 1404 wolves were caught
(Brauner, 1923). In the eighties of the past century, there were many
wolves in Chernigov governance (Dubyanskii, 1885).

At the end of the nineteenth century, the number of wolves in the
entire middle zone and in the Ukraine noticeably decreased. At the
beginning of the twentieth century the wolves remained in significant
numbers only in districts with large tracts of forests and sparse popu-
lations. In [the former] Khersonsk, Nikolaevsk and Odessa provinces,
they were completely destroyed, and in the other southern districts
wolves remained only in the floodplain of the Danube (Izmail’sk dis-
trict). At the time of the First World War, the number of wolves grew
sharply; this was enhanced by the Civil War and the devastation ac-
companying it. By 1920, the wolves occupied nearly all of the Ukraine,
central chernozem district, the middle non-chernozem zone and the
Povolzh’e. In Chernigov district in 1923 alone, wolves destroyed 5.2%
of the domestic livestock count. As was noted, wolves were, at that
time, very numerous in the north of Kustanaisk district (V.G. Heptner).

In subsequent times, the number of wolves began to decrease and
at the beginning of the Great Patriotic War [W.W. II] wolves de-
creased overall by approximately 100 times as compared to 1920. At
this time in the Ukraine, wolves remained only in Zhitomirsk, the
southern regions of Kievsk and Chernigovsk, and southwestern regions
of Khar’kovsk districts, in the Starobel’sk steppe and in Lugansk dis-
trict (Migulin, 1938; Korneev, 1950).

New proliferation and recolonization of wolves took place in the
years of the Patriotic War, 1941-1945. The number of wolves grew
intensively in places where in addition to food abundance, wolves had
well protected places for constructing dens. After the Great Patriotic War,
the number of wolves increased in all districts without exception, and
they appeared in the places where they had been previously exterminated.
In the Caucasus, where the number of wolves had increased somewhat

after 1927—1930 and remained at quite a high level until the Second World
War, it sharply increased in the period 1941-1945. Only after 1950, as а
result of the strengthening of wolf control did their numbers in the Caucasus
noticeably decrease (Vereshchagin, 1959).

Thus also proceeded [events] in the European north and in western
and eastern Siberia. The distribution of wolves in Siberia was favored
by winters with little snow at the end of the 1940’s. Wolves became
the scourge of [rein]deer breeding in the Katangsk region of eastern
Siberia and appeared in the Shitkinsk and Bodaibinsk regions (Timofeev,
1949).

After the war, wolf control was sharply intensified, and their num-
ber quickly began to decrease everywhere. This was favored by the
application of new, totally destructive methods of hunting (see below).
In Arkhangel’sk district, intensive destruction of wolves stopped their
growth in numbers, and then sharply reduced them. The number of
captures per year in Arkhangel’sk district increased from 129 in 1948
to 360 in 1953 (in all 1374 wolves were destroyed in 1948—1953;
Semenov, 1954).

In addition to this basic factor in the population dynamics of wolves,
great significance must be placed on nutritional conditions in winter
and the supply of water in summer. The first is of basic importance in
the forest zone and the second, in the steppe and desert zones. How-
ever, in the steppe regions of Kazakhstan, the number of wolves also
sharply decreases in winters with much snow, mainly as a result of
death from starvation. In the winter of 1948/49, which was character-
ized by strong and prolonged blizzards, wolves were not able to capture
food. They gathered near the settled points and somehow fed on dogs,
but they quickly began to die (Afanas’ev et al., 1953). At the same
time, the widespread opinion that all wolves are obliged to fast in
winter is not true. In a pack (family) of two grown-up wolves and four
juveniles caught in December 1940 in Moscovsk district (near
Maloyaroslavts), the adults were well nourished, but not fat, while the
young had good deposits of internal fat which nearly covered the kidneys
and a little fat on the thighs. It is evident that the old wolves, even in winter,
were solicitous about the better nutrition of the young. Accessible livestock
are not here in winter, and only carrion was at the disposal of the pack,
apparently, elk, hares and dogs being scarce in the district (in the stomach
of one wolf, a piece of leather collar was found; Heptner and Morozova-
Turova, 1951). Communications about the capture, in winter, of fat and
“very fat” wolves are not very rare, particularly from the tundra
(Makridin, 1959).

258

Fig. 70. Tracks of a pacing yearling female. Oka preserve, Ryazansk district.
15 January 1949. Photograph by V.G. Heptner.

After a series of drought years in semi-desert and desert regions,
the number of wolves sharply decreases. Thus, in 1939 in the Ustyurt
and Mangyshlak, wolves almost totally disappeared, and in the Turgaisk
region of Kustaniask district in 1938, the number of predators sharply

186 decreased. The drying-out there of steppe springs and lakes deprived
the wolves of waterholes and the surviving animals concentrated in
abundance in ravines at the village of Borgot. In the bottom of ravines,
the wolves dug holes, in which ground water gathered, and the animals
came regularly to water at these “excavations” (Afanas’ev et al., 1953).

259

In the drought of 1936, all wolf litters living in the lower Sarysu river
and adjacent regions, gathered in small reeds surrounding the two re-
maining Telekul’ lakes. They attacked everything available for them to
pursue, but they apparently were starving and the young grew poorly
(Sludski, 1937).

Among wolves epizootics sometimes develop, but their causes,
course and after-effects are practically unstudied.

Fig. 71. Tracks of pacing wolf. Krasnoyarsk territory. December 1959
Photograph by V.V. Kozlov.

187

260

The reproduction of wolves in different years is not the same. In the
central chernozem districts, intensive reproduction was observed in 1925,
1934 and 1935, in 1943, 1944 and 1947 (Barabash-Nikiforov, 1947).

Field characteristics. The wolf is a strong, enduring animal; with-
out special effort it can pull or carry on its back a ram. It can move
quickly, achieving a speed on bare ground* of 50—60 km per hour
(Semenov, 1954). Information about speeds up to 80 km per hour
(Nanteifel’, 1948) is completely unverified. Usually wolves run at a
trot. This trot nonetheless allows them to cover great distances. In this
gait, the track of the hind foot falls exactly on the track of the fore, and
the tracks are situated, as along a string in one line. During walking,
this exactness in the position of the feet is not observed. Wolves of a
well-fed pack go in single file, and a tired lead animal is replaced
periodically. This allows the pack to move for long distances even in
deep snow. Hungry wolves sometimes separate from each other.

When running, the wolf holds its head somewhat low and a little
to the side, directing one ear forward and the other backward. This pose
gives them the possibility of using their exceptionally sharp hearing.
Changing the gait when necessary, the animals switch to a gallop and
then a walk; they often stop and stamp in place. In deep snow, they
prefer to use roads. By day they go at a walk, avoiding populated
places; sometimes dodging and doubling back, like a hare, usually
shortly before lying down. Alarmed, wolves may go for 70 km in one
night at a trot or jog (Korneev, 1950). Wolves can swim well. In
autumn 1948, a litter of wolves regularly swam across the Desna river,
on setting out to hunt and on returning from it to the den until a strong
cold snap [occurred] (Korneev, 1950). In Ryazansk district in the 1950’s,
in periods of night hunting, they swam across the swift, wide Oka river
to attack the livestock on an island (V.G. Heptner). It is very difficult
to catch up with untired animals by rowboat.

The tracks of wolves resemble those of dogs, but differ in their
form and large size. The wolf does not “spread” the digits in the track,
but holds the pads, as hunters say, more “balled up”. The print of the
anterior pads of a large central Russian wolf reaches 123 mm in length
and 55 mm in breadth. The track of one wolf in the Priozern region of
Arkhangel’sk district had a length of 130 mm and a breadth of 97 mm.
The track of a huge animal, killed in Vladimirsk district in 1953, was
15 x 12 cm in shallow freshly fallen snow (N.D. Sysoev). The largest

*Literally chernotropa, or “black path;’ see р. 242—Sci. Ed.

261

track of the anterior pad of a tundra wolf is 126 x 92 cm, and of the
posterior, 123 x 80 mm (Timansk tundra; Semenov, 1954). The soles
of the pads of the tundra wolf are better furred and the toes are more
movable (expanding more widely). In tracks, females and males can be
differentiated according to the manner of urination (the first is directly
on the tracks, the second to the side). The she-wolf and the young leave
their feces directly on the tracks, whereas the wolf usually moves aside
from the tracks.

Trails of wolves differ from individual tracks in the somewhat
larger size of the tracks, their compactness, the wider “drag”—the
furrow in the snow, inscribed by the end of the wolf pad in the begin-
ning of the step, and the “pillow case’”—such a furrow at the end of
the step (in front of the foot print).

Fig. 72. Wolf trail. Track of four wolves proceeding “track in track”. Khakassiya,
Krasnoyarsk territory. December 1956. Photograph by V.V. Kozlov.

188

189

262

The voice of a mature wolf is a long bass howling. The male wolf
gives voice through an octave, passing to a deep bass with stress on
“О”, and the she-wolf produces a modulated baritone through the nose,
with stress оп “О”. Her voice is particularly mournful and wild. Yearlings
howl with characteristic yelps resembling those of dogs. They include
high notes and end by a characteristic barking. Juveniles almost never
howl, but whine with laugh-like, thin voices. Howling wolves are heard
for a distance of up to 2 km. The meaning of the howl is an assembly
call for the mature wolves returning with prey and a signal of well-
being in the region of the den. The male howls first, the female answers,
and the young repeat in a dissonant chorus. Howling begins at twilight
after sunset, and continues with pauses of usually not more than 15
minutes. Wolves do not answer a howl in rainy weather and when
satiated (Danilov, 1945). In autumn and spring (until the birth of young),
and sometimes in winter, wolves usually howl when going to hunt. In
certain periods, for example, in the forests of the lower Samur river in
southern Dagestan in August 1924, they “struck up a tune” with great
precision at the coming of darkness (V.G. Heptner) (N.N.).

Practical Significance

Wolves are harmful to humans in many respects: they attack livestock
and dogs, wild ungulates and other useful animals, spread diseases and
attack people directly. The little use which may be derived from cap-
tured wolves (skin, tasty meat which is fully suitable for food) as well
as sporting pleasure of the hunting of wolves, are not to be compared
with the damage to human health and economic interests caused by
these undoubtedly injurious predators.

The average agricultural losses caused by wolves in the middle of
the past century in only the 45 European governances of Russia were
estimated at 179 thousand head of large horned livestock and 563
thousand of small annually, for a value of 7 million 600 thousand
rubles in those day’s prices (Lazarevskii, 1976). Animal husbandry in
Vologodsk, Novgorodsk, Minsk, Volynsk, Kaluzhsk, Orlovsk, Ryazansk
and Samarsk governances suffered especially from wolves. In 1871—
1873, wolves killed an average of 2550 (0.29%) head of large horned
livestock and 8550 (0.65%) of small livestock per year in Kiev gov-
ernance alone. In Volynsk governance at that time, mortality reached
1.36% per year (12,300 head of large horned livestock and 19,600 of
small; Korneev, 1950). Recently, the amount of harm sometimes

263

Cr Lo
Wot 9S ace "<= ofc ec = э= 9
ove
3 Fo Ce
eet al eines о ое 5
Dog (4 ar
1
Fea
Wolf x x “= =
LA
р v= we ste =
Dog ore 9
C=
“= = be
Wolf Ка = é =
CS eo
“> > xe <=
=> Cu. © ve =. Su
Wolf = aS 4 x =

188 Fig. 73. Scheme of tracks of wolf and dog in different paces of running (from
Kozlov, 1955). 1—walking slowly, 2—trotting, 3—pacing, 4—zalloping.

188 Fig. 74. Prints of front and hind feet of wolf on soft silt. Ganyshkino, in the delta
of the Volga. 23 October 1951. Sketch by A.N. Formozov, about 1/2 natural size.

264

decreased and sometimes increased, growing particularly during social
and economic shocks (wars, economic crises, crop failure and so on)
as a result of weakening of predator control. This was noticed both with
us, and abroad as well.

The harm caused by wolves greatly increased due to growth of
their number after the First World War and then in the Civil War. In
three republics: the Russian Federation, the Ukraine, and Uzbekistan—
865 thousand head of livestock were registered as killed by predators
in the agricultural year 1924/25, a sum of 15,530 thousand rubles in
the price at that time. Approximately 35% of the losses took place in
Siberia and Kazakhstan, followed by the other main stock-raising dis-
tricts. In relation to all livestock in general, these losses consisted, in
the lower Volga of 2.2%, in Siberia—1.6%, in Kazakhstan—1.5%,
and for the whole territory of the USSR, 0.54% on average. Among the
animals killed, 16% were horses (4.2% adults, 11.8% young), 18.1%—
large horned livestock (6.8% adults, 11.3% young), 52.6%—sheep,
5.8%—goats and 7.2%—swine (Krasil’nikov, 1926). About 30% of
the animals were killed in winter in the time of confinement in struc-
tures or in winter quarters. Many domestic poultry and dogs are killed
by wolves. Hundreds of thousands of geese were exterminated yearly.
In summer, they sometimes even comprised the principal food of wolves
in a series of places. Although losses were reduced in 1928 due to the
return of wolf control, until 1941 they remained in the vicinity of
several hundred thousand head of livestock annually, 1.е., quite high.

In connection with growth in the number of wolves at the time of
the Great Patriotic War and afterward, there was a sharp increase in
damage caused by them. Thus, in 1944 in the Buryat ASSR, more than
5300 head of livestock were destroyed by wolves; in Penzensk district,
780 horses and cattle, more than 5000 sheep, 300 swine and others
were killed in 10 months of that year. Losses for the whole year
approximated 8700 head of livestock in all districts. In Kuibyshevsk
district, this figure reached 4200 head of livestock. In sheep-raising

‚ state farms, the wolves killed at that time 1—1.5% of the breeding
flocks each year (Danilov, 1945). In Vornezh district, only the wolf
population connected with Usmansk forest caused losses in the forest
itself, and on 295 collective farms situated in its neighborhood, esti-
mated at 760 thousand rubles in four years (1946—1949). During that
time wolves killed on collective farms 164 calves, 163 foals, 3119
goats and sheep, 37 pigs, 3174 dogs, and 2500 geese. At that time as

190 well, in Usmansk forest wolves destroyed 200 deer and not less than

265

19 beavers (Martts, 1953; Barabash-Nikiforov, 1957). In 1946 in the
Zyryanovsk and Asinovsk regions of Tomsk district, 600 head of live-
stock were slaughtered by wolves (Boikov, 1949).

The amount of damage caused by one litter of wolves can be
judged by the fact that in Ryazansk district (Bel’kovsk region) 2 ma-
ture and 6 young wolves destroyed during 8 months (from May to
December 1945) 6 horses, 1 foal, 5 cows, 150 sheep and goats, 3 pigs
and many domestic geese, ducks and hens (Kozlov, 1955).

Annual losses to reindeer-breeding caused by wolves in the Yamalo-
Nenetsk Nats. Region amounted to 1% of the population of the deer
herds; in 1948, losses reached 2% (Rakhmanin, 1959). In 1951 they
killed 1708, and scattered 7048 deer which “disappeared without a
trace”. In a decade (1944—1954) these losses amounted to about 75,000
deer (Semenov, 1954). In 1952 in Arkhangel’sk district, wolves and
bears killed 1282 horses, 1343 head of large horned livestock, and
1806 sheep and goats. Of these losses, 64% took place in 7 southern
regions of the district where wolves were especially numerous. Even
greater damage to reindeer breeding was caused by wolves. Mortality
of 3—7% of the domestic reindeer population was from wolves (Korsh,
1936). Losses of livestock due to wolves in Azerbaidzhan comprised
20-25% of the total losses in livestock breeding after the war
(Vereshchagin, 1957).

These examples show how much the great losses after 1941—
1945 were caused by the multiplication of wolves. To this must also
be added the damage which is inflicted by wolves on dog husbandry
by the destruction of hunting dogs during hunts, sometimes nearly in
front of the hunter’s eyes. Laikas and hounds in particular suffer from
wolves, which follow their voices. However, wolves also attack watch-
dogs, falling on them right in villages, or luring them beyond the
outskirts.

Wolves cause great harm to the hunting economy, killing hares,
foxes, game birds and waterfowl, wild ungulates, as well as muskrats,
raccoon-dogs and even sables (Koryakov, 1947). Elimination of young
wild pigs, red deer, roe deer, chamois and Kuban turs by wolves in the
Caucasian preserve consisted in only 5 months of 34 to 61% (Teplov,
1938). In Il’mensk preserve, a litter of wolves in 180 days killed about
100 roe deer, or 15 animals per month on the average (S. Severtsov,
1941). In the lower Ili river, half of the roe deer and wild pigs born
are killed by wolves in autumn. In years with deep snow, wolves may
kill all of the young (Afanas’ev et al., 1953). In the winter of deep

191

266

snow in 1940/41 in Bashkir preserve, 35 roe deer were found killed by
wolves, and in the winter of 1942/43, the remains of 8 bull and cow
marals were met with (not all, of course, were found). A family of 7
wolves ate a whole bull maral during one night, leaving only shreds of
the hide (Kirikov, 1952).

One wolf crossing ice to Barsa-Kel’mes island (Aral Sea) slaugh-
tered 34 goitered gazelles and | saiga from March to June. Moreover,
beginning in May, it ate the eggs and nestlings of gulls and caught sand
susliks [Spermophilus fulvus] (Afanas’ev et al., 1953). Wolves cause
particular harm to wild ungulates in the steppe and desert after calving,
when they kill the young en masse (Heptner, 1956).

The intensification of wolf control in the post-war years sharply
decreased the “wolf threat”. However, it still continues to demand
serious attention. Even though there is a relatively small number of
predators, they are dangerous due to their boldness and bloodthirsti-
ness. Attacking a rich quarry, for example a flock of sheep, the wolves
kill much more than they need in order to eat. In December 1949, a
wolf slaughtered 60 geese in one night on a poultry farm in Zhitomirsk
district (Korneev, 1950). A litter of wolves, especially at the time of
“hunting training” can kill 60—80 sheep in one flock (Sokolov, 1951).

In the autumn of 1920, on the headwaters of the Tobol in Kustanaisk
district, wolves (probably a litter) on a dark night attacked a flock
spending the night on the steppe, separated it from the shepherd and
drove off the frightened sheep in the darkness, killing one after the
other. In the morning, 52 carcasses were found on the steppe, lying in
a long line in the direction of the flock’s flight. Not far away, wolves
attacked a herd of horses and slaughtered about 10 foals. All the
animals were killed with a characteristic wolf bite—their throats were
torn (V.G. Heptner). A wolf got into a sheepyard in winter in Redkovichi
village (Belorussia) and killed 18 sheep in one night (Serzhanin, 1955).
In June 1941 in the Izhevsk region of Ryazansk district, 2 mature
wolves attacked a flock of sheep in a corral and in the course of one
hour slaughtered 96 animals. On the night of | January 1950 in the
Shilovsk region of Ryazansk district, 4 wolves got into a collective
farm sheepfold and killed 78 sheep (Kozlov, 1955). At the end of June
1957 in Kopanov village in Ryazansk district, a wolf jumped. over a
wattle fence almost 2 meters high, and burst into a pen near the sheepfold
where the sheep were driven at night. It killed and wounded 55 sheep,
of which 5 in all were treatable. There was no watchman and the
animal worked unhindered. Apparently, it did not carry off anything.

192

Fig. 75. Remains of saiga killed by a pair of wolves at a waterhole. Chu river
valley, Kazakhstan. 3 November 1955. Photograph by A.A. Sludskii.

After this attack the pen was raised 70 cm and barbed wire stretched.
After several days, the wolf reappeared and again penetrated the pen
through a narrow space in the sheepfold wall where the barbed wire
was not stretched. This time there were watchmen who raised an alarm,
thereby giving the animal the possibility of a fortunate exit. It wounded
12 head in all of which 3 could be treated. In this way, the wolf
destroyed in all 59 head in 2 nights (V.G. Heptner).

Wolves are the most dangerous vectors of rabies, not only because
dogs and other animals (and recently, raccoon-dogs) are infected through
them, but also because the bite of a rabid wolf is, in its consequences,
about 15 times more dangerous than the rabid dog (Rudnev, 1950).
Moreover, wolves serve as important vectors of helminths for domestic
animals. The role of wolves in the transmission of the causative agent
of “sheep gid[diness]” (Multiceps multiceps), and in part echinococ-
cus, is of great importance.

Finally, even healthy wolves, although in very rare cases it is true,
attack humans. More or less verified cases of attacks by healthy (not
rabid) wolves on people have been registered in Belorussia (Serzhanin,

268

1955), in Kirovsk and Irkutsk districts (Timofeev, 1949), in the Kareliya
(Marvin, 1959), and in the Ukraine (Korneev, 1950). All of them took
place in winter, the time in which wolves are hungry.

Thus, in December 1947 in the vicinity of Suna station on the
Kirov railway, a large man-eating wolf prowled about for one month.
It usually went out around the village in the morning and evening,
caught dogs and attacked solitary people, in daytime lying in the bushes
near the road, not far from the village. In the course of one month, this
wolf attacked people about 15 times. It bit and scratched 13 people, and
tore to death two others (a woman and an adolescent). Killed after
some time this wolf was found to be a large old male (head—-body
length 138 cm) that was extraordinarily emaciated. Except for a bundle
of woman’s hair, nothing was found in its stomach. Apparently, this
wolf became a man-eater, being unable to catch ungulates. This was
aided by the fact that during the war, eating of corpses by wolves was
a frequent occurrence (Marvin, 1959).

In Ukraine, cases have been described when wolves attacked women
in Dnepropetrovsk district in the winter of 1923/24 (Barabash-Nikiforov,
1928); two cases were registered in Stanislavsk district, one in
Chernigovsk (attack on a shepherd in August), one—in Kamenets-
Podol’sk and one—in Poltavsk district, when wolves killed three children
(Korneev, 1950).

Control of wolves is carried out in various places by different
means. Universally distributed are: a) extermination of litters in spring
and summer in dens, when the pups are not yet grown and do not
participate in hunting with their parents; b) capture of wolves with
traps, especially in winter; c) application of poisoned baits. Traps are
placed in trails (“under the track” or “in the track”), by lures, on scent
trails, and near the so-called “urination points” (see above). Besides
poisons which kill the animals (strichnine and others), soporific means
(for example luminal) have now begun to be used by placing them in
baits.

In the forest and forest-steppe zones in the second half of summer,
autumn, and winter, they are hunted with an enclosure or by driving
with or without application of flags. Hunting wolves with guns (“when
howling’’) is successful. Near baits, and in summer also near the den,
they stand watch with their guns. In the forest and forest steppe, bor-
zois and hound dogs are used; in the east they are hunted with golden
eagles, and sometimes the hunters bring wolves to bay on horseback.
This method was particularly widespread in Kazakhstan. For this were

193

269

required the best (“baigov’)* horses, although wolves do not have
great endurance when galloping. At full gallop, the animal is killed by

-a blow to the head with a heavy whip (kamchi).

For this hunting the Ural and Orenburg Kazakhs employed a long
stick with a root-like knob on the kom/*. On such an “ukryuk” [lasso],
a loop is usually fixed for catching horses from the herd. This method,
to some extent, takes on a sporting character; however, capturing in this
way was in the past one of the chief, if not the only method in the
steppe, and in part, in the forest-steppe habitats of Kazakhstan (V.G.
Heptner).

Self-snaring of wolves (loops, frames, cages, pits) are not efficient
and have local significance; in the Buryatsk ASSR through self-snar-
ing, 60 wolves were caught in 1946 (Manteifel’ and Larin, 1949). In
Armenia, wolves are caught in winter in the unfrozen irrigation canals
with quick current and cement walls by building a false bridge (Dal’,
1954).

The significance of these methods of exterminating wolves is not
equal. According to data derived from a competition on the best means
of extermination of wolves in 1927, the largest number of predators
was caught by thorough extermination of litters in dens in summer
(9.6—5.6 wolves per hunter); in second place was control by poisons
(7.3-4.7 wolves per hunter); third—driving on horses (5.5 wolves per
hunter); fourth—capturing with traps (4-3 wolves per hunter); fifth—
capturing with dogs (3 wolves per hunter); sixth—by shooting from
ambush near carrion (from 2.7 to 4.7 wolves); and seventh—drives
using small flags (from 1.4 to 5.2 wolves) (Fedosov, 1939; Danilov,
1945).

In 1946, 263 wolf cubs were destroyed in dens in the steppe
regions of Ryazansk district alone, in Voronezh—265, and 637 cubs in
Astrakhansk. One hunter in Kirovsk district in the summer of 1950
exterminated 38 wolves by shooting them “when howling”; another got
17 wolves in this manner in 1949 and in 1950—36 wolves. With a
golden eagle, one hunter caught in 1938/39, 154 foxes and 9 wolves,
and another, in 1952/1953, caught 124 foxes and 26 wolves.

During the last few years, in open landscapes (tundra, forest-tun-
dra, forest-steppe, steppe, desert) and even in the forest zone, the method
of extermination from airplanes and airsled is found widely applied. In
the winter of 1945/46 in Kuibyshevsk district, one pilot cornered and

*Turkic (?) word—Sci. Ed.

270

ow”

Ne Eg Е ЕС ВЕРА ВЕТ a
ОТ HAGE SUR SS НД MAS о СОЯ ОО
1936 1940 1950

Fig. 76. Graph of tanned wolf hides as an indication of change in wolf numbers
in the Caucasus by year. Continuous line—Cis-Caucasus; broken line—Trans-
Caucasus (from Vereshchagin, 1959).

destroyed 180 wolves. Two pilots in the winter of 1947/48 killed 264
wolves from a PO-2 airplane in Tatariya and Chuvashiya (Popov and
Lukin, 1949); in Kalininsk district, in the same winter, 200 wolves
were caught from a PO-2 airplane (Kozlov, 1955). In March 1949, in
Volgogradsk district, 38 wolves were killed from two airplanes in 7
days. In the Nenetsk Nats. Region, up to 1955, the average yearly
number of wolves captured by all methods was 57 head (including
cubs), a fact that restrained the growth in the number of wolves but did
not decrease it. After employment of airplanes, the number of captured
wolves grew sharply and the number of wolves and damage from them
were reduced. Thus in 1955, 92 predators were killed by airplanes
while all land-based hunters caught only 72 wolves; in 1956, 72 wolves
were taken from airplanes, and 48 by land-based hunters (Makridin,
1959).

The wolves are very successfully driven and killed by airsled. It is
applied in combination with an airplane (for reconnaissance) and airsled
in radio contact sent by orders from the airplane for destruction of the
wolf packs thus found (N.N).

194

271
Genus of Arctic Foxes
Genus Alopex Kaup, 1829

1829. Alopex. Kaup, Skizz d. europ. Thierwelt, I, p. 83, 85. Canis
lagopus Linnaeus. .

1868. Leucocyon. Gray. Proc. Zool. Soc. London, p. 521. Canis lagopus
Linnaeus (У.Н.).

Forms of moderate size.

Skull fairly compact with relatively narrowly placed zygomatic
arches, quite massive and moderately high (height of the occipital re-
gion comprises about 1/3 the condylobasal length). Sculpturing of skull,
particularly sagittal crest, weakly developed, but greater than in g[enus]
Vulpes. Facial portions moderately elongated (relatively shorter than in
foxes of g[enus] Vulpes), quite massive but weaker than in the repre-
sentatives of g[enus] Canis. Distance from line connecting the ends of
supraorbital processes to anterior end of premaxillae longer than that
to upper edge of occipital crest. Interorbital (frontal) area convex,
somewhat elevated in area of frontal sinuses and, on the whole, notice-
ably and quite abruptly elevated above line of upper rostral profile.
Supraorbital processes quite weak, flat or even slightly concave from
above with tapering tip which is directed slightly downwards. Posterior
edge of tooth row extends to level of anterior border of interpterygoidal
notch. Deep indentation and protuberance on lower edge of lower jaw
in front of angular process absent.

3 1 + 2

Dental formula igs ox aps Maen 42. Teeth comparatively
moderate in size and massiveness, carnassial well developed; length of
upper carnassial and cheek teeth goes into palatal length about 2%
times. Canines quite massive, markedly curved and of moderate length
(when the jaw 1$ closed, tips of upper canines extend to the middle depth
of mandible, while tips of lower canines reach alveoli of upper). Tips
of upper canines directed straight downward; no distinct longitudinal
cutting ridge on posterior side of canine and lateral surface of canine
smooth. Incisors have no supplemental cusps, or they are very weakly
developed.

Body notably elongated, tail quite long—about half the body length,
ending considerably below the tarsal joint, and in standing animals reaching
the ground. Head elongated. Ears long with rounded tip. On the soles of the

196

272

hind feet there are only 4 digital pads—behind them there is no large fifth
pad.

The fur is very dense, luxuriant, long and soft. Winter fur is
monotone and dimorphic—white, or more rarely grayish-brown; sum-
mer fur is dark in color with light flecks. There are two molts per year.

Supracaudal* gland not developed. From 5 to 8 pairs of teats (A.F.
Shirkova)'. Evident sexual dimorphism is absent; age is well marked,
and seasonal dimorphism is very sharp, not only in the degree of
density and length of fur but also in color.

Forms of the genus possess a quite typical foxy appearance.

They are inhabitants of the tundra and the Arctic islands. They
accomplish regular seasonal migrations. They are monogamous, uniting
together in the reproductive season. Up to 18 offspring (perhaps up to
20—22—the maximal number in the family and order), but usually less.
The young are born only in a burrow. They are predators, feeding on
birds and small mammals (rodents), flotsam, and carrion. Apart from
the breeding period, they pursue a solitary way of life.

The range of the genus is remarkable in that it consists of two
parts. One of these represents the region of reproduction and is occu-
pied by the animals in summer time; the other is an area of transience
and wintering, which they occupy in winter. The region of reproduction
occupies the whole zone of tundra and the Arctic islands in the Old as
well as the New World. The region of wintering and winter migrations
extends on the north to the ice of the Arctic Ocean, and on the south
to the region of stunted trees [krummholz] and the northern part of the
forest zone’.

Morphologically, the genus is somewhat less specialized than Vulpes.
Physiologically, as typical arctic [forms], this group is very sharply
specialized. The genus stands as a quite well-defined intermediate group
between the genera Canis (wolves) and Vulpes (foxes). In some re-
spects, it is closer to Vulpes and has been considered only a subgenus
of the latter, although it has enough characters that ally it to the genus
Canis also. More than anything, on the basis of the “intermediate”

**Violaceous” in Russian original—Sci. Ed.

'Reference to 6 (3 pairs) of mammae in the arctic fox, widespread in foreign
literature, is incorrect. Such a number of таттае is not possible in an animal which
gives birth to more than 10—12 young.

*Mammals migrating in a definite season from a more or less considerable part of
their breeding range are very few. Cases in which the species leaves almost all its range
are exceptional for the terrestrial mammals if some bats are not considered. Such type
of migration is quite analogous to the migration of some birds.

‘IouldayH ‘9`Л ‘э8иел рэ}эп1$и05э
“URDDO, INDY dy) JO 9d! dy} UO Чиой oY} 0} зиоцел8 ци JOJUIM JO зцопоэир итеш эч—5моме :ррол\ мэм OY} Ul YINOS oy) 0} зиоцел8 ни JURYSIP
Арепоцие@ yenpiarpur jo saoejd эзеотрит syulog ‘лэзитм ит иоцел8 ни Jo Алерипод Woyynos—oul] иэхол4 ‘(8итииэр) ээиэ11950 JoWIUNS JO зэмерипоа
—эи| рцоз :рром PIO oy) ш "71 $7408] ха4о]у хо} эполе эц} JO э8ие1 $91994$ oy) рие dney ха4о]у ‘эхо; эполе JO $пиэ8 ay} JO эЗиел OY] “LL “BI4 S61

a. %
te
{=

(ег)
os 09 05

og:

274

characteristics of the genus Alopex, one may unite all three genera into
one and consider them as subgenera. It has been assumed that Alopex
is a subgenus of this broader genus Canis, which includes other groups
besides the three already named (see above, characteristics of the
family and of the genus Canis). At present, however, the majority of
the authors consider the group А/орех as an independent genus. Until
there is a more detailed analysis of the family system as a whole, this
point of view is accepted here.

The genus is known from the Pleistocene (Europe), but it was,
apparently, established in the upper Pliocene.

During the period of narrow interpretation of the species, some
species were accepted into the genus (some authors considered up to 5
of them). It is now certain that the genus comprises only one species
which constitutes about 3.5% of the species of the family.

In the fauna of the USSR, there is one species, Alopex lagopus
Linnaeus, 1758; constituting 0.3% of the total species number of the
fauna.

Its distribution in the USSR covers the tundra and northern islands
from the Kola peninsula to Cape Chelyuskin and the Commander
Islands.

This genus is one of the most important fur-bearing species in the
USSR (V.H.).

ARCTIC FOX [POLAR FOX, PESETS]
Alopex lagopus Linnaeus, 1758

1758. Canis lagopus Linnaeus. Syst. Nat., Ed. X, 1, p. 40. Lapland.

1898. Alopex lagopus spitzbergenensis. Barret-Hamilton et Bonhote.
Ann. Mag. Nat. Hist., 1, p. 287. Spitsbergen.

1902. Alopex lagopus beringensis. Merriam. Proc. Zool. Soc. Wash.,
15, p. 171. Bering Island, Commander Islands.

1912. Vulpes lagopus beringianus’. Suvorov. Commander Islands and
fur industry on them. p. 217. Commander Islands.

1931. Alopex beringensis semenovi. Ognev. Zveri Vost. Evropy i Sev.
Azii [Animals of Eastern Europe and Northern Asia], vol. 11,
p. 265. Mednyi Island, Commander Islands (V.H.).

’This transcription is sometimes wrongly ascribed to Cherskii, 1920 (Ellerman
and Morrison Scott, 1951).

197

275
Diagnosis
The only species of the genus.
Description

In general appearance, the arctic fox is very similar to the common
fox (Vulpes vulpes). In its short summer fur, it appears to have relatively
longer limbs than the fox, with a shorter body. In its luxuriant winter fur,
it seems to be more squat, although not so extended. The head of the animal
in its winter fur is fairly rounded, with a somewhat blunt face and very
widely separated ears which appear slightly above the fur; the ear tips are
rounded. In summer fur, the head looks disproportionately large, with large
and widely separated ears. The tail is Jong, about half the body length and,
in winter fur is covered with long and luxuriant, extraordinarily dense hairs.
When the animal is standing, the tail reaches the ground. The soles of the
feet are covered with dense and long hairs, foot pads are not visible.
The summer fur 15. dense, soft and delicate, but short. The general
tone on the back and neck is smoky-grayish-brown, sometimes with a
mixture of reddish tones, that on the side is the same but with a lighter

‘tone which passes to whitish on the belly. The thighs have the color of

the sides and the legs, including the feet, are covered with brownish-
gray fur. The dark color of the back in the region of the shoulder blades
descends like a cape to the limbs and thus forms a cross-shaped figure

Fig. 78. Arctic foxes, Alopex lagopus L.—white (left) and blue. Sketch by
A.N. Komarov.

198

276

(the summer Arctic fox is called “cross-fox’’). The inner parts of the
limbs are brighter than the outer. The tail above is brownish-gray, and
the remainder is lighter, mixed with some pale-reddish-brown tints. The
head is brownish-gray, the dorsal side of the ears is gray-brown and
their front and inner parts are white.

The winter fur is extraordinarily long, dense, luxuriant and
soft. The long and soft guard hairs form a beautiful “veil” of fur, and
the very firm and dense underfur are characteristic. The color is
entirely snow white, only sometimes with hardly noticeable creamy
tints. In addition to this normal white color phase characteristic of the
great majority of the animals throughout the entire range with the
exception of some insignificant points, there is also a dark phase, called
“blue fox”. The color of animals of this phase is very variable
and varies from sandy, gray and light coffee-color to dark ash-gray
with bluish tint; light and dark chestnut also occur, often with a more
or less well distinct silveriness (white guard hairs). The whole body is
of one color. The summer fur of arctic foxes of the dark phase does
not differ, or only slightly differs, from the winter. The summer fur is
of one tone and the “krestovatik’ [“cross”] pattern* does not occur.

Sexual differences in color are absent, while age-related ones are
quite considerable. New-borns are covered with short velvety fur of
dark-brown color, only very slightly lighter on the sides. With age the
lightening of color on the sides becomes stronger and gradually form
the figure in the summer fur described above, and the little one leaves
the burrow in a “young krestovatik” pattern*.

For characteristics of the skull and dental system, see above in
description of the genus.

The diploid number of chromosomes is 54.

The size of arctic foxes is subject to quite considerable individual
variation. Moreover, the average size of individuals of one population
may vary annually. Extreme and average weights of the animals of one
and the same population in different years may differ very greatly,
depending on food reserves which annually change very sharply, per-
haps being entirely different. Besides, the nutrient status of animals
differs at different times in winter—the highest weight is in the begin-
ning of winter, and by spring it strongly declines. In a good year,
weight may increase even up to December, and its decrease begins with
January. There is also geographic variation in size.

*See above—Sci. Ed.

Fig. 79. Blue fox, Bering Island, October 1958. Photograph by S.V. Marakov.

Body length, 46.5—73 cm, tail length 25—52 cm. Weight changes
from 2 to 8.8 kg (combined data of mainland and island animals,
including Commander arctic foxes, but without Spitsbergen; see be-
yond, “Geographic Variation”). In the Lena delta in winter
(November—March) 1935/36, weight of males (118) was 2400—6800 g,
an average of 3816 +* 109 g; that of females (114) was 1200—5600
g, average 3091 + 107 g; for the whole population the extreme variants
(232) were 1200—6800 g, average 3453 + 55 g. In Table 19, changes
in weight in various months are given (ТаугоузКи, 1939).

Greatest length of skull in males is 127—144.3 mm; in females,
119—136 mm; condylobasal length of skull in males is 114—138.5;
in females, 111.0—127.2; zygomatic width in males is 64.7—78 mm;
in females, 63.2—77.0 mm; length of upper tooth row in males is

*Omitted in Russian original—Sci. Ed.

278

` ``,
BN
= ,

Ominnee

199 Fig. 80. Skull of arctic fox, Alopex lagopus L.

200

279

53.1-61.3 mm; in females, 50.4—59 mm (skull dimensions include агс-
tic foxes of the Commander Islands, but exclude Spitsbergen) (V.H.).

Table 19. Weight of foxes in the Lena delta by months (in g[rams})

Males? Females
Months SSE
Minimum Maximum Average Minimum Maximum Average
November 2400 5600 3753 1200 4600 3129
December 2700 5400 4036 2220 5600 3500
January 2500 6800 4012 2000 4400 2989
February 3000 4400 3607 2400 3700 2988
March 2450 3200 2850 1900 3000 2350

Systematic Position
The only species of the genus.
Geographic Distribution

The extreme north of Eurasia and North America. The borders of the
range change considerably seasonally—the region of reproduction oc-
cupies the tundra and the Arctic islands, in the period of winter migration,
it covers the northern part of the forest zone and the ice of the southern
Arctic ocean. The region of irregular winter migrations of individual
animals extends still farther, both to the north and to the south.

Geographic Range in the Soviet Union

This comprises a great part of the range of the species. Within the
borders of our country is to be found in nearly all the Eurasian region
of occurrence of the species. It occupies a relatively small part of the
State territory on its northern frontier.

The southern boundary of the region of summer occurrence and
regular reproduction in general follows fairly closely the southern
boundary of the tundra, but in places, this strict coincidence is some-
what disturbed. On the Kola peninsula, the above-mentioned boundary

ЗИ is significant that through January inclusively, the amplitude of weight fluctua-
tions is very high—the smallest individuals are more than two times lighter than the
heaviest. In the second half of winter, this contrast is less. Apparently, this takes place,
to a certain extent, as a result of the decrease in the difference between young and old
animals at the end of the winter (V.H.).

280

proceeds from Varanger Fiord, a little north of Murmansk, to the mouth
of the Ponoi on the eastern shore of the peninsula. In places, the arctic
foxes, under the suitable conditions (montane tundra), den somewhat to
the south of this line.

On the eastern coast of the White Sea, the border passes a little to
the north of the mouth of the Mezen’, extends across Malozemel’sk
tundra, mainly its maritime parts, and the northern part of the Timansk
mountain chain. Crossing the extreme lower Pechora, the border goes
farther to the east, along the watershed and the upper reaches of the
Bol’shezemel’sk tundra rivers, 1.e. along its middle and southern parts.
At the Urals, the border extends nearly to the Arctic Circle or a bit
north of it. On the Urals, the border descends to the south along the
montane tundras.*

To the east, the boundary line slightly ascends to the north, reach-
ing the mouths of Shchuch’ and Ob’, encircling Salekhard to the north
and reaches the mouths of Nadym and Pur. Along the divide of the Pur
and Taz, it descends quite a considerable distance to the south to the
Arctic Circle. Farther to the east, the border passes northward along the
right bank of the Taz beyond the Arctic Circle, crosses the Yenisei at
the level of Noril’sk lake, includes them, and along the mountainous
places (Putorana mountains?), makes an extension southward, includ-
ing the sources of the Kureika and Kotue, and apparently Lake Essei,
not reaching south, however, to the Arctic Circle.

From the above-mentioned place, the boundary rises quite abruptly
to the north, crossing the Khatanga in its lower course and proceeding
to the north of 70° N. Lat., crossing the Lena at its lowest reach.
Farther on, it slightly descends towards the south, crossing at about 70°
the Yana and Indigirka, and extending south of 70°, reaches the mouth
of the Kolyma in the area of Nizhne-Kolymsk. From here, this line
extends to the southeast, reaches the upper Anadyr’, and thence abruptly
turns to the south and even somewhat to the southwest towards
Gizhiginsk gulf, thus becoming the western boundary for Chukotsko-
Koryatsk part of the range. The southern border here, i.e. in the direction
of Kamchatka, passes nearly along 60°, cutting across the very north-
ern part of the peninsula. This is the southernmost known point of
occurrence (denning) of the arctic fox.

With respect to information concerning the occurrence of the arctic
fox in Kamchatka, they are contradictory. According to old data
(Krasheninnikov, 1762 and others), the arctic fox was encountered in

*Not evident on map, Fig. 81—Sci. Ed.

281

the entire peninsula or along its eastern coast to Cape Ukinsk (not far
from Cape Kamchatka), and along the western coast—to its southern
extremity (Cape Lopatka) or somewhat to its north. This information
was interpreted by some authors as permanent occurrence, and by
others, as winter migrations. In recent time there appeared a commu-
nication (Lavrov and Naumov, 1948), that a permanently occupied
colony of arctic foxes is on the coast of the Sea of Okhotsk in the Ust’-
Khairyuzov region. All of this information is, for our time, either old
or doubtful. One may think that the arctic fox is not present now in
Kamchatka or it is extremely rare. In the past, it is apparent that, on
the peninsula in the above-mentioned areas, transient animals (regu-
larly migrating), were caught and farther south there were only rare
irregular transgressions.

The northern border of the continental part of the range is every-
where formed by the ocean. Moreover, arctic foxes den in all the Arctic
islands from Kolguev and Franz Josef Land to Wrangel’ Island, and
also on Kil’din on the Murmansk coast and on Karagin and Com-
mander islands in the Bering Sea. Arctic foxes are absent only on
several very small islands, lying in the open ocean, such as De Long
Island. However, arctic foxes were found on Uedineniya Island in the
Kara Sea (V.G. Heptner).

In spite of some old assertions, aboriginal arctic foxes are absent
on the Kuril islands. In 1915—1916, blue foxes were brought from the
Commander Islands to Ushishir (Usisira) Island where they survived
well and reproduced. In the 20’s and 30’s, arctic foxes were brought
to the islands of Simushir (Sinsiru), and from the small Kuril islands
of Yuru (Yuri, Uriri), Kharukaru* and Lis’i (?, V.H.)**. On the Malyi
[small] Kuril islands at the present time (60’s) foxes are apparently
absent (Kuznetsov, 1939; V.G. Voronov and A.G. Voronov).

The southern border of the area of regular migrations—the winter
range—in the extreme west passes along the southern coast of the Kola
peninsula. Within Kareliya, normal migrations are not common, and
here only occasional long movements are known. The southern border
of the above-mentioned area between the White Sea and the Urals
traverses the northern part of the forest zone (V.I. Parovshchikov), i.e.
somewhat south of the Arctic Circle, although not rarely the arctic fox
also penetrates far to the south.

*Kharimkotan?—Sci. Ed.

**This query by Heptner is not explained, but may refer to the source(s) from
which the above data are derived. In particular, the context of the sentence suggests that
foxes were not brought from, but rather, to the small Kuril islands—Sci. Ed.

49
e 30
170

}
ВИН
но 160 160

CN Е
$ =
}

0 1 130

а
NS
а и at

203

202

283

To the east, the boundary of winter habitation crosses the Urals
and the Ob’ somewhat to the south of 65°, reaching the upper Pur and
Taz, or in the vicinity of their watersheds with Vakh, and crosses the
Yenisei at Verkhne-Imbatsk (below the mouth of the Podkamennaya
Tungus). Thereafter it goes to the source of the Vilyui and crosses the
Lena, Yana and Indigirka rivers around the Arctic Circle or a bit more
to the north, passes over the Kolyma slightly above Verkhne-Kolymsk
(considerably south of the Arctic Circle) and, still descending south-
ward, reaches the Penzhinsk Gulf. This district includes the northern
and western (excepting the extreme south) parts of Kamchatka.

In general, it may be assumed that winter migrations of the arctic
fox extend to the northern parts of the forest zone—sometimes quite
deep into it. In such cases, apparently foxes of the extreme west (Kola
peninsula) roam to the south least of all; there is comparatively little
shifting of the range in the remaining European part of the [Soviet]
Union and in Yakutiya, and animals in west and central Siberia move
southward farthest of all. Evidently very distant migrations occur in
places in northeastern Siberia and Kamchatka.

Migration of separate individuals does not always take place di-
rectly south- or northward. Many animals also move in other directions.
In different regions, there may be different predominant directions.
Thus, arctic foxes from Yamal move mainly to the southwest and
straight west, going as far as Cheshsk Gulf. Animals during these
journeys put behind them many hundreds, even more than a thousand,
kilometers, without even leaving the region of summer distribution of
the species. This probably explains the fact that in the European part
of the [Soviet] Union, the border of the winter habitation of the arctic
fox shifts comparatively little (for more see description of migrations
in the section “Biology”).

Fig. 81. The range of the arctic fox, Alopex lagopus L., in the USSR
(reconstructed). The continuous line limits the southern region of summer
habitation and reproduction (denning), the dashed line—the region of regular
wintering and permanent winter migrations. Dots indicate places of most distant
winter transgressions of separate individuals to the south, stars—to similar places
to the north on the ice of the Arctic Ocean, figures near them give the latitude.
The question mark in Kamchatka refers to the place (Ust’-Khairyuzovo) regarding
which there is information about the denning of a small group of animals in an
isolated spot, triangles—places of Upper Pleistocene finds of arctic fox. V.G.
Heptner.

284

The outlined southern boundary of the region of normal winter
distribution is entirely tentative. The reason for this is above all an
insufficiency of accurate materials. Moreover, this line is much less
sharp in nature than the boundary of the area of denning. Depending
on various reasons, in one and the same district, the distance of
migration to the south and the number of animals coming from the
north, may annually change noticeably. Therefore, the border may be
situated in the south sometimes far and sometimes near, while at the
same time, in various parts of the range, these annual variations may
be expressed in various degrees. Finally, in a series of cases, with the
presence of data in the literature, it may be difficult to decide whether,
in a given place, we deal with regular residence of a small number
of individuals, i.e. with the district of normal wintering, or with a
place whence separate vagrant individuals migrate comparatively
frequently, i.e. with a district of long-distance transgressions (in the
present description, the southern boundary of the normal migration to
the south is directed, apparently, somewhat more to the south than is
accepted by some authors, at least for separate parts of the range).

To the south of the region of regular wintering is situated a wide
zone of irregular transgression of separate vagrant individuals. They
often transgress so far to the south, that they, apparently, do not have
the possibility of returning back, and in the long run they probably
die. The following most distant places of transgression are known
(extreme points, from west to east, Fig. 81; for more see the section
“Biology”): Baltic Sea coast at Ventspils (Vindavy; Van den Brink,
1958), Daugavpils on the Zapadnaya Dvina (about 56° N. Lat.), up-
per Velikaya river, flowing into Pskovsk lake, Velikie Luki region,
northwest of Yaroslavl’ (58° N. Lat.), northwest of Kirov at 60° N.
Lat., the Omutninsk region in the upper Viatka at 58°40’ М. Lat.,
northwest of Tobol’sk, approximately at 59° N. Lat., Yalutorovsk on
the Tobol at 56°40’ М. Lat., middle course of the Vasyugan—a tribu-
tary of the Ob’, the mouth of the Parabel’ at the Ob’, Kolpashovo on
the Ob’ at 58°30’ М. Lat., the Chulym river to the north of Achink
(about 57° N. Lat.), Kansk (west of Krasnoyarsk), Taishet (west of
Krasnoyarsk at 56° N. Lat.), Bratsk on the Angara, Cheremkhovo on
the Angara (to the northwest of Irkutsk at approximately 53° N. Lat.);
one of the southernmost points in Siberia), Bodaibo on the Vitim
(about 58° N. Lat.), Mukhtui on the Lena (about 60° N. Lat.),
Olekminsk on the Lena, Yakutsk,.mouth of the Tompo river at the
Aldan, source of the Tompo river (about 64° N. Lat.), Oimyakon

204

285

(about 63° N. Lat.). Tauisk Gulf on the Sea of Okhotsk, southern
Kamchatka (about 52° N. Lat.), northern Kurils. Migrations to the
lower Amur and Komsomol’sk-on-Amur—S50° М. Lat. are known
(points from personal material of A.F. Chirkova). It is possible that
migrations to the lower Amur may be in part connected with the
movement of animals along the ice or drifting with floating ice.
Therefore, in the European part of the country migrations extend to
56° N. Lat., in the Asiatic—nearly to 50° N. Lat. This difference
could be explained by the fact that conditions characteristic of arctic
fox are, in Siberia, distributed farther to the south than in the west;
in particular, montane tundra extends far to the south. Another reason
is that human settlement in the west is much more dense and the
animals die earlier, before they attain their natural movement.*

Winter outposts of arctic fox from the region of reproduction
occur not only in the south, but also in the north—on the sea ice.
Movement in this direction represents an entirely normal phenom-
enon, since in its wandering on the ice, the arctic fox usually follows
the [polar] bear. It is evident that in southern latitudes and districts
of the Arctic Ocean, this is a “regular” winter habitation to very
distant places—“irregular long-distance transgressions.” However, the
definition of these districts is still almost impossible.

As shown by observations in recent years, including both our
own and also American drifting research stations, arctic foxes are
more common in the region of the southern sea margin and southern
parts of the Arctic Ocean, and the farther toward the polar district, the
rarer they become. Known extreme points of arctic fox finds show
that they penetrate to the most inaccessible parts of the Arctic Ocean
farthest from land, and that they probably reach the pole. Very many
cases of arctic fox observations are known from the ice region rela-
tively close to the continent (up to 100—150 km). The following are
the most northerly encounters in the waters in our sector of the
Arctic: 77°32' М. Lat—132°59' Е. Long.; 78°19’ М. Lat—141°33’ Е.
Long.; 78°27’ М. Lat—177° Е. Long.°; 78°30’ М. Lat—176°03' Е.
Long.; 78°42' М. Lat—152°33’ Е. Long.; 80°30’ М. Lat—159°50’ W.
Long.; 80°51’ М. Lat—176°00' W. Long.; 85°20’ М. Lat.—about

“Information concerning the capture of an arctic fox in the Tien Shan, which is
widely found in the literature (for example, Ognev, 1931), and even used for some
historical formulations is based on a misunderstanding: the skin pertains to an albino
red fox (Afanas’ev et al., 1953).

*“РгауЧа” newspaper оп 16 April 1941.

205

286

60° Е. Long. (our northernmost point; Chapskii, 1946; Rutilevskii,
and Uspenskii, 1957).

All points mentioned are distant from the nearest land by several
hundred kilometres—in a biological sense, this is more important
than the latitude. Thus, the place of occurrence at 85°20’ М. Lat.

-stands 550 km from Rudolf Island (Franz Josef Land); the point at

78°27’ Lat. lies more than 500 km distant from De Long Island and
more than 750 km from Wrangel Island; the point at 78°30’ lies 750
km from the New Siberia[n Islands]; that at 80°51'—900 km from
land; and the point 80°30’ and 159°50' W. Long. found in the “pole
of inaccessibility”, is 1025 km from Point Barrow in Alaska and 1125
km from Wrangel Island. In the American sector of the Arctic, the
arctic fox was noted at 87°, 460 km north of the northern tip of
Greenland, and at 88° N. Lat., 800 km from Ellesmere Land.

In the Quaternary period, the arctic fox was distributed much
farther to the south. Its Pleistocene remains are known in all countries
of Western and Central Europe, including Ireland, southern France
(Pyrenees), Switzerland, Czechoslovakia, and Poland. Within our
boundaries, the remains of arctic foxes are known from the Gomel’,
Bryansk, Lipetsk, Chernigov, Poltava, Dnepropetrovsk regions, and
even the Crimea. In the east, Pleistocene arctic foxes were found in
the Chelyabinsk, Krasnoyarsk and Nizhneudinsk regions. Therefore,
they could be found in the south as far as 44°40’ М. Lat. (Crimea)
and about 55° N. Lat. (Siberia). This demonstrates that contemporary
transgressions of arctic fox, both in the European part of our country,
and in the east (Pri-Baikaliya) reach or nearly reach the places of
their Quaternary occurrence.°

It is remarkable that in spite of very ancient and very intensive
exploitation, not considering the comparatively small territory of
Kamchatka, the range of the arctic fox did not suffer any essential
changes.

SRange according to data of Krasheninnikov, 1786; Maak, 1859; Middendorf,
1887; Pleske, 1887; Ognev, 1926, 1931; Solov’ev, 1927; V.N. Skalon, 1928; Formozov,
1927; Paramonov, 1929; Yakovlev, 1930; S.P. Naumov, 1931; N.P. Naumov, 1934;
Tugarinov, Smirnov and Ivanov, 1934; Kolyushev, 1936; Heptner, 1937; Sdobnikov,
1940; Chapskii, 1946; Kuznetsov, 1949; S.P. Naumov and Lavrov, 1948; Stroganov,
1949; Shvarts, Pavlinin and Danilov, 1951; Yanushevich and Blagoveshchenskii, 1952;
Lable, 1953; N.I. Dement’ev, 1955 and others, and according to unpublished materials
of V.N. Skalon, V.Ya. Parovshchikov and V.G. Heptner.

For additional data concerning transgressions, see section on “Biology”.

287
Geographic Range outside the Soviet Union

In Europe—the extreme north of Scandinavia and the highest parts of
the mountains of the peninsula facing the Atlantic Ocean, as a narrow
strip to the south, nearly to 60° N. Lat. In winter wanderings—all of
Finland, and all Norway and Sweden to its extreme south. Appar-
ently, they migrate onto the ice of the Baltic Sea. The islands of
Spitsbergen, Jan Mayan, and Iceland.

In North America, the southern boundary of the range covers the
northern parts of Labrador, the Ungava peninsula, and the district
lying to the east of Hudson Bay. Across the bay and beginning on its
western shore slightly south of 60° N. Lat., the boundary passes
westward approximately along this parallel, south around Great Slave
Lake and along the Mackenzie river to its delta. Farther on, it encom-
passes the northern and western parts of Alaska, reaching the ocean
at Cook Inlet. To the north of this line, the range includes the entire
northern periphery of the continent and all the islands of the Ameri-
can Arctic and all adjacent districts of Greeniand. In the Bering Sea,
the range includes the Aleutian and Pribilof Islands and Hall, Saint
Matthew and Saint Lawrence islands. On winter migration, arctic
foxes reach as far south as the Gulf of Saint Lawrence, and the
northern parts of Alberta, Saskachewan and Manitoba (i.e. approxi-
mately 55° N. Lat.). Particularly distant transgressions extend to 50°
and even 45° N. Lat. (Nova Scotia). (V.H.).

Geographic Variation

Compared to the ranges of such species as the wolf or red fox, the
range of the arctic fox is not great’. Moreover, it is distributed in one
landscape-geographic zone, and in the various parts of the range
there are no sharp contrasts in conditions of existence which may
stimulate geographic variation as an adaptive process. Finally, the
arctic fox is characterized by seasonal migrations, since some animals
move for enormous distances, without leaving the region of habita-
tion and reproduction of the species. This provides a high level of
panmixia in the species population. Thanks to the continuous cover
of arctic ice and the wide migrations of animals on the ice, island

’The exaggerated extent the dimensions of the range of the arctic fox is based
on the wide use of maps with Mercator projection.

206

288

isolation is practically not met with. Within the boundaries of our
country, it exists only in relation to the Commander islands, where a
separate race was formed. In the parts of the range outside the USSR,
the same conditions and the same tendency towards the formation of
forms are generally revealed.

In our part of the range, the following subspecies are distin-
guished.

1. Continental arctic fox. A. [. lagopus Linnaeus, 1758.

Color of winter fur is snowy-white, of summer fur smoky-brown-
ish-gray (see above); the dark phase is very rare and is usually
encountered in the proportion | : 1000—2000. Changes in this ratio in
different parts of the range are negligible. Only on Kil’din Island оп
Murmansk coast, the percentage of blue reached 30—50.

Skull? of moderate dimensions, quite broad and high; muzzle
relatively long (its length constitutes 43-48% of condylobasal length);
interorbital region wide; infraorbital processes of moderate size;
postorbital constriction not sharp; braincase quite high and wide—its
height represents 36-43%, and width—3441% of condylobasal length.
Cranial crests not large, but well defined. Proximal ends of nasal
bones located at the level of the upper borders of the frontal pro-
cesses of the maxillary bones or somewhat beneath it, rarely higher.

Size not large: body length 45—70 cm, weight to 7 kg.

Greatest length of skull of males 127—136 mm, of females 119—
131 mm: condylobasal length of skull of males 114-132 mm, of
females 111-127 mm: zygomatic breadth of males 64.7—73.7, of
females 63.2-70.3: length of upper tooth row of males 54.1-—61.3, of
females 50.4—59.0.

Continental part of the range, from the Kola peninsula to Chukotka
land: all Arctic islands except, perhaps, Franz Josef Land.

Not recorded outside the borders of the USSR, but perhaps dis-
tributed in Alaska.

The Novaya Zemlya arctic fox were, in part provisionally, some-
times separated as A. |. spitzbergenesis. This, however, is unfounded,
for they cannot be differentiated from those of the continent by skull
dimensions. The above-mentioned form is characterized by very small
size (condylobasal length of skull 105—120 mm), and infantile skull
type—the skull of adult Spitsbergen animals corresponds to that of

*Characteristics of the skull here and later after Tsalkin (1944). Dimensions of
skull after Ognev (1931).

289

subadult continental ones. That the arctic foxes of the Franz Josef
archipelago are related, or close to, the Spitsbergen form, is not
excluded, however, nothing is known of them.

Craniologically, arctic foxes found in the whole of the above-
mentioned expanse are identical; however, fur merchants differentiate
here the following types (Kuznetsov, 1952).

Novaya Zemlya type. Skins of this type are differentiated by
especially high quality. They are characterized by large dimensions
(weight of 100 skins about 30 kg), luxurious silky pelage with tall,
elastic guard hair and dense underfur, and by the snow-white color.
Flesh side of skin compact.

Novaya Zemlya.

Yenisei type. Skin quality nearly equal to Novaya Zemlya and
almost the same dimensions (100 hides weigh 28 kg). Pelage only
slightly less silky.

Tundra between Taz and Anabara.

Yakutsk type. Skin dimensions slightly less than the former (weight
of 100 hides 25.5 kg). Pelage characterized by fluffy, and chiefly, by
exceptional silkiness; guard hair fine, delicate, less elastic and under-
fur dense. Color clear-white. Flesh side of skin thin.

Yakutsk tundra.

Obdorsk type. Skins of moderate size (weight of 100 hides less
than 27 kg). Pelage not so fluffy and markedly less silky than that of
preceding types, with short, especially on the back, elastic guard hair

Fig. 82. Ratio of fur types in tanned skins of white arctic fox in tundras of

Arkhangel’sk district in the year of movement of arctic fox from the east (of 100%

at each point) (А.Е. Chirkova): 1—Pechora type, characteristic of European
North; 2—Obdorsk type, characteristic of Yamal.

290

and dense underfur. Color white, but usually with light creamy tone.
Flesh side of skin compact.

Region between the Urals and Taz.

Pechora type. Skin dimensions small (100 hides weigh 26 kg).
Pelage less fluffy, with short guard hair and loose underfur. Color
white, with barely noticeable bluish tone (color of skimmed milk).
Flesh side of skin compact.

Tundras of the European part of [Soviet] Union.

2. Beringian arctic fox, A. [. beringensis Merriam, 1902.

Color of winter fur light or dark-brownish, usually with more or
less well-defined silvery highlights. Dark smoky skins with deep blu-
ish underfur (“dark blue”) are relatively scarce. Fur is extremely
fluffy, but somewhat coarse, with well-developed guard hair. White
color phase is absent or found in the form of extremely rare indi-
vidual cases.”

Compared to skull of continental arctic fox, skull narrower in
region of zygoma and braincase, its height less, supraorbital pro-
cesses and postorbital constriction more sharply defined, teeth larger,
crests more strongly developed.

Measurements are somewhat larger than continental ones. Body
length of males 46—67, average 60 cm, females 41—64, average 57 ст;
weight of males 15 3.5-8.8, average 5.5 kg, females 2.5—6.4, average,
4.4 kg.

Condylobasal length of skull of males 121—139 mm, females
116-134 mm. Length of nasal bones in males 41-57 mm, females
41-50 mm.

Bering Island.

Absent outside the borders of USSR.

3. Mednyi [Copper Island] arctic fox, A. /. semenovi Ognev, 1931.

Color of winter fur of the majority of animals is dark-brownish
without admixture of white and silvery hairs. Clear-gray (“blue”)
arctic foxes occur in comparatively small numbers. Fur is somewhat
coarse, with weak, scattered guard hair. White color phase is absent
or is found in the form of extremely rare individual cases.'°

°Characteristics of color and fur, and general dimensions of this and following
forms mainly after Kuznetsov (1952).

In the past, with suppression of the predominate dark color phase, as on both
Mednyi island and on Bering island, white arctic foxes were met with more often than
in the present time. The present situation is the result of the deliberate elimination of
the white forms over the course of many decades.

208

291

Skull noticeably larger than that of continental arctic fox, narrow
in zygoma and in region of braincase, its height less, nasal bones
shorter, supraorbital processes and postorbital constriction more sharply
defined, teeth stronger. Skull crests, especially lambdoidal and region
of interparietal bone are very robust—much more robust than in
continental animals.

Dimensions considerably larger than in continental arctic fox,
and somewhat larger than in Beringian one. Body length of males 61—
73, average 68 cm, females 60—68, average 64 cm. Weight of males
4-8.8, average 7.2 kg, females 4—7.2, average 6 kg.

Greatest length of skull of males 130—144.3, females 122.6—130;
condylobasal length of skull of males 116—139 mm, females 116—131
mm; zygomatic width of males 72—78 mm, females 66—77 mm; length
of nasal bones of males 36-45 mm, females 34—44 mm.

Mednyi [Copper] Island.

Absent outside the boundaries of USSR.

Bering and Mednyi arctic foxes are extremely close to one an-
other, although the differences between them are quite real. Mednovsk
are also distinguished from Beringian in averaging shorter nasal bones.

It is worth mentioning that, at the same time that European island
races of arctic foxes (spitzbergenensis, fuliginosus) are quite regu-
larly characterized by their small measurements and by some infantile
features of skull structure, in Commander (Aleutian and Pribilof)
arctic foxes, these are not observed. They, on the contrary, are larger
than continental ones, with more specialized skulls (with features of
hypermorphosis). Probably, this is explained by the very favorable
conditions of existence, in the sense of food supply, of arctic foxes
on the above-mentioned islands.

For the part of the range lying outside our boundaries, the follow-
ing subspecies are usually accepted.

1) A. JL. spitzbergenensis—Barr.-Ham. et Bonh., 1898—
Spitzbergen; 2) A. [. fuliginosus Bechst., 1799—lceland; 3) A. 1.
groenlandicus Bechst., 1799—Greenland; 4) A. [. ungava Merr.,
1884—Labrador and Ungava; 5) A. [. innuitus Merr., 1902-—Conti-
nental tundras and islands of America, except the regions occupied
by the forms ungava and groenlandicus; 6) А. |. pribilofensis Merr.,
1902—Pribilof islands; 7) A. [. halensis Merr., 1900—Hall, Saint
Lawrence and Saint Matthew islands in the Bering Sea.

Arctic foxes of the American islands—Pribilof and Aleutian, like
the Commander islands, represent the dark phase. Apparently, not all

292

the listed forms are sufficiently well characterized and well grounded .
(УВ:

Biology

Population. In the tundra zone of the USSR, the arctic fox is the most
common and abundant carnivore, substituting here for the red fox.

The number of arctic fox is extremely unstable; it sharply changes
by years and by seasons of the year. In autumn and at the beginning
of winter, in some years, one may encounter tens and even more
arctic foxes in the course of a day, and in other years, especially in
spring, only a single individual in the course of a month.

For orientation, one may assume that the overall arctic fox popu-
lation in the season before the beginning of harvest amounts, in the
Soviet sector of the tundra zoné, in different years, to from several
tens of thousands to several hundreds of thousands''. Arctic foxes are
most abundant in the Bol’shezemel’sk, Yamal’sk, Prieniseisk, Prilensk
and Ust’yansk tundras.

Arctic fox stocks are not great in the narrow belt of the Iokan’ go-
Ponoisk tundra on the Kola peninsula (see Table 20 and beyond). In
the tundras of northern Arkhangel’sk district, the arctic fox inhabits
most densely the northeastern Bol’shezemel’sk tundra. In some years,
these tundras are full of arctic foxes migrating from the east. Arctic
fox is quite abundant in the north of the Kanin peninsula (Table 21).
In a year of small numbers of arctic foxes (winter 1951/52) in an area
of 400 km/?, only 3 arctic foxes were taken in a drive; in a year of
high numbers (winter 1953/54) in several drives on an area of 20
km’, about 15—30 arctic foxes were taken in December, and in the
second half of winter about 7—15 arctic foxes.

In the Malozemel’sk and Bol’shezemel’sk tundras during the
time of land development work, the general number of burrows was
revealed (Table 21), the density of which in the northern subzone of
the tundras increased 3 times. The total number of arctic foxes was
arrived at (Table 20) by an average “yield”, by considering 30% of
the holes as being occupied and by taking into account two pups for
each breeding. In these tundras, 7 large “hearths” of reproduction of

"'Boitsov (1937), taking the area of tundra as 3,116,500 km’, considered the
number of arctic foxes in the 30’s to be 100,000 breeding pairs. From this, each pair
of animals inhabits, on the average, 31 km? or a population density equal to 1.3 per
1000 ha.

209

293

arctic foxes were discovered, with the density of burrows ranging
from 2.9 to 5.6 per 1000 ha in each, and an average of 4.8. The area
of the hearths comprises about 16% of the entire land of Malozemel’sk
and Bol’shezemel’sk tundras, and includes about 27% of all discov-
ered burrows (Skrobov, 1958). In the east of the Bol’shezemel’sk
tundra, burrows are located at a distance of 1—2 km from each other,
and rarely at a distance of 400—500 m (А.Т. Voilochnikov, Yu.T.
Nazarenko). At the mouth of the Pechora, in autumn 1944, with
abundance of arctic foxes close to the multiyear average, hunters
found in one harvest day 3—10 polar foxes, while in autumn 1954,
with numbers negligible, 1-2 foxes in 10 days.

In winter 1956/57, on a route of more than 500 km in the region
of Kara Bay at the extreme northeast of Bol’shezemel’sk tundra, for
each 20 km of the journey, from 19 tracks in November to | track
in January, averaging 2—6, were found. In winter 1957/58 in a year
of diminished numbers of arctic foxes was recorded (on a route
length 1500 km) an average of 0.7 tracks (Chirkova, 1955; Chirkova
et al., 1959). The fox population in Novaya Zemlya was calculated
to be several thousand head (Dubrovskii, 1937) (Table 20).

In Yamal, a great number of arctic foxes live in the northern
subzones of the tundra (Dubrovskii, 1940; Freiberg, 1940;
Tsetsevinskii, 1940; Chirkova, 1955; Kopein, 1959; N.N. Spitsyn),
but, apparently, not in the extreme northern edge of the peninsula
(Table 21), where the tundras exist on a very flat plain. Along the
Venui-euo, Tiutei, Kharosovoi, Se-yaga, Morade and Tambei rivers,
burrows of arctic foxes are particularly abundant. At places in the
middle course of the Tiutei river, burrows are located at a distance
of only some tens of meters from each other. The region of massive
denning of arctic foxes in the maritime tundra of the southwestern
coast of Baidaratsk Bay connects with the best arctic fox lands on the
Bol’shezemel’sk tundra. Some authors consider that in 1/2 of the
entire Yamal territory, burrows are few, and probably only in 1/4 of
the territory are they met with frequently. It is suggested that in
Yamal, there are about 10 thousand burrows of arctic foxes; this
constitutes 1/3 compared to the Bol’shezemel’sk and Malozemel’sk
tundras. Judging by the abundance and catches of arctic foxes, it is
evident that the number of burrows here is considerably greater (see
the first footnote to Table 21).

Already at the end of the last century, Nosilov (1909) who wan-
dered in Yamal for more than 2 thousand km, could see only 6 arctic

"ззиел8ти 34ээхЗь,
‘PISIA э8елэле шол с,
‘SoBe ПУ,

м—

0'S-0'7 8561 {2561 24 001 UO [661 ‘AOSHOG {6561 ‘OYUOyHLL
0001~ :008- 8261 ‘S761 7561 риа! oy) Jo Алонлэз omNUg ‘9561 ‘B1aqio14 ‘рие$1 Мирэи

ЕС: 1 8061 ‘ey 001 YO
00S °€~ :000°7~ 761 pues! oy) Jo Алоэ} эзциЯ 6761 ‘OYUSYHIL :9561 ‘ЗлэатэлА
рие!$1 Sullog ‘5рие|$] Jopuewwos
ЕЕ Е ee ee ee PES CUNO роб tepueWwWUO)
St C470 6961-9961 ‘ey 0001 UO 6$61 ‘AOJOZO]Ig pue ло3$14$

‘JOLSIP uepesep|
ЕЕ Е a a ee a Se en

€ 1-10 Алепие (1 эашэлом ‘661 Ч 0001 YO 6561 ‘91941214
1$. JO UOISAI ‘лэлы BUD] 40 2

Be 19913456 *L 46] Ч 0001 YO 8561 ‘лохтиЧорб
чэлы ваАцие | eAvAuYZIN ‘еазитэа JAWIe

| 9661 Ч 0001 YO [561 “AOSHOY

[ITN 004-69 ^9|э4 :A10}1LI0} узиечуил1,
a ea о МО 07 ISN ees eee

000°8 ueRYI э10 | 9561—5561 10314133 [561 “HysAosqngd

000°r~ 61-61 peyqeyur эти ‘eA[WaZ вАвлоМ

ЕЕ ЕВЕ Е Е С Е АМ

СС 80 Ч 0001 YO 8061 ‘^040435 ‘5влрип}

00S SI $961 561 AlOVIIO} SIU — BABYS JOWOZO[eI рие вАеу5.[2щ197245.109

—— es oe ee Ne eee
ald ‘sired © 03 Jowiuns *7¢6] ‘ey 000! YO

aves ‘eu OOSL 1661 ‘AOSHOG ‘6561 ‘AOZOWIO

‘sited пре 0у—66 UeY} элош 10N Jaquiojdag-jsn3ny ‘/ 76] ALOWIII} IU ‘pueys] UIP [Py

АЕ Е ee ee se Se ee ee
vIpUN) YSIOUDg-03 ,ULYO]

0007-0001 ALO) OUT ‘e[Nnsuludd е10У

a a ek eo ee ee Se ИВА
$эхо} 18104 Jo Jaquiny Чоп] JO OWI вэгу лоЧуие ‘и018э$
ee rere oe nee et tO

suonendod хо} 3199.18 Jo UOTE] NI]ed ay} Jo ajqu) PauIquIOD “OZ aIqGU

605

‘моли рэмаечи!г АЦеэтромэ4 Jo Jaquinu |014,
‘SMOLING YOT'ZZ INOge “(€961) AOGOIYS 0} Зирлоээу,

м—

50 рэмаечиий «f9YZOYN,, OC JO vole ue Uj 9161-5161 L761 ‘^лололп$

70 Рэи9ечи ‘pueys] Suliog
a ee ea
Jot]1e9 0561—6161 “HPISIOyD

$/- рэиаечч] QUOZ [815805 эЦ} Jo Алоулл |, рие 8161 рие[$1 'Кирэи{ ‘зрие|$1 лэривииио

$'0—$0'0 еЧ 0001 YO 7615561 [761 ‘лоцешоз ‘у

9:000°9 SeIpUN} ay} JO AOI] oINUT = = puke 1 76|—-9Z6| Алоэ eSULIeYY-CUudT
eee eee Ee oe St
(5 wo) (Zw 00S 176] ‘Aourwioy ‘у

0's JO э9в1эле JO Bale [101 WO) „мя OO] UT 0761 SIOALI 41519 Jo you

Jind у$8иезецу 40 15809 usojsea ‘ejnsutusg JAWNIE]

PAOMNYD “AV

81 гу | 5.061 *(е4!4о 109

67 uo $99819 ul Sey 0001 UO ‘OUIZNSIB A “ВАвлэишо) лэлы тэзшэ д. JOMO'T
Е ee
81 ‘BY 0001 YO 5.0761 pues] чо5жа
а eee
70 JO э8е1эле ‘ey 0001 YO 9761 LE6I “AOSHOG “387 °М .0L-69

(11 wo) | 2W 06 1эло э8елэлу м0|э4 Ал10)114э3 YsueyyNIn] jo seipuny
Е бе
000`01-000°6 seipun} oy} Jo А10}14123 эзиЯ jewex 1 pue ejnsuiusg ysuepAH
Е ee A ee ee
Z—-1< JO э8е1эле ‘ey 0001 YO veol—Ce6l 0761 “HP{SUIAaSIOS L

51000'01 Ало31ллэ3 aU 1561 ‘uXsuds ‘N'N

‘азии [ешел,

—з ыы —[—щ—=—=—=—=ы—=—=—=—=—=—=—=—=—д——————————————.—.—.—.—.—..—.—.—._.._._.„.„„ЗЗЗАЗЭЗЗ ences

ZTE JO э8елэле

0'S-8'1 SOUOZQNS SNOHeA JO BY (0001 YO 8561 “AOGOLIS

000‘0€ 1 stipun} oy) Jo ArOjI9} элция SS6I-£S61 ‘SBIPUN} у$,[2щ1220]е/\{ рие yS,jawWezoys [09
И а
(SMOlING 0001 Wo.) “ey 0001 чо Lv6I-Sr6l 1561 “Aoueqiys
8°S-L'0 ‘ejnsuluag uluey

Е о, a Pi ee ee ee SE a ee ee SS НЯ
змолиа JO JaquINN вэту 1х 1оЧуие ‘и018э

м—ввмыььъБъБъЪъЪчнтньнкБ д ——а—а—

SMOLING XOJ 319918 JO повете» Jo э[4 Е) paulqwiosy *[Z э14ЕТ,

OI?

296

foxes, but the Nenety [native people} told him that “arctic fox was
found not a long time ago and it was encountered on each verst”. In
our time in northern Yamal, in the summer, in a year of average
“vield” (1949), the number of arctic foxes met with during one day’s
journey by sled* is 1-2, and in winter up to 10 tracks were encoun-
tered; in the Pri-Ural region in August, up to 10 arctic foxes and in
winter 2—5 animals. In the Mare-Sale region on the western coast of
Yamal in autumn, they used to wander in tens, and only after the
freezing period did their numbers sharply decline. On the coast of
Baidaratsk bay, in the middle of August 1947, in a year of abundance,
on a walk of 80-100 km, more than a hundred arctic foxes were
counted and in winter, at places for | km, there were 1—2 arctic foxes
and 3—5 tracks. Deep in the tundra, arctic foxes were significantly
fewer; on a journey extending for 80—100 km, only 10 foxes were
met with. With an abundance of animals in southern Yamal in the
winter of 1947/48, the hunters caught from 15 to 35 arctic foxes for
the season, and in the best conditions, up to 70. In a drive on an area
of 20—30 km’, 40—60 arctic foxes were taken in December—January
1955/56 and in February—March, 20-30. In the years when their
numbers are depressed, on a week’s excursion in the tundra, 2—3
tracks were encountered and only isolated arctic foxes were caught
in the season; it was only the best hunters in southern Yamal who
were able to take 10—15 individuals (A.F. Chirkova).

The total number of arctic foxes in the Yamal region after the
end of the harvest season for 1955—1958 was 3.1—3.6 thousand head—
1/4—1/5 of those, that were in the hunting period (12.4—17.6 thousand
per year). The degree of harvesting was 70-80% of the population
(Smirnov, 1959). However, judging by the number of arctic foxes
hunted in this region, in which a considerable part of the territory was
still insufficiently developed for harvest, one may presume that the
number is considerably greater, especially at the beginning of winter.

In the northern part of Gydansk peninsula and in the Malyi
Yamal peninsula, along the Poilovo and Le-Yakha rivers, a large
concentration of arctic fox burrows is observed. In the territory of the
latter 21 sections with abundant burrows were revealed by an aero-
plane. The number of burrows in the Gydansk peninsula and Malyi
Yamal is assumed to be equal to the number in Bol’shoi Yamal
(Table 21). But, considering the extent of territory of the Gydansk

*Russian word is narta, which means both dog- or reindeer sled—Sci. Ed.

212

297

peninsula, the sparse settlement and poorly developed harvest of arctic
foxes, a significantly greater number of burrows may be expected
here.

In the north of Krasnoyarsk territory, arctic fox is comparatively
densely settled in the tundra of the lower Yenisei, along Pyasina river
and in part the northeastern Taimyr peninsula (Rutilevsku, 1939;
Shastin, 1939; A. Romanov, 1941; Chirkova, 1955; Tables 20 and
21). In the Avamsk tundra, burrows in places are encountered at a
distance of 10 km from each other. In northern Yakutiya, the western
regions are richer in arctic foxes than the eastern ones. There are
many arctic foxes on the New Siberian and Lyakhovsk islands and
between the Lena and Indigirka [rivers] (Konstantinov, 1921;
Tugarinov, Зтитоу and Ivanov, 1934; Freiberg, 1939; A. Romanov,
1941).

In the Lena-Khatanga territory, the density of burrows declines
from north to south. In summer, the maritime tundra between the
Khatanga and Anabara, between Anabara and Olenek north of the
Pronchishchev range, and between the Olenek and Lena in the region
of the Chekanovsk range are densely inhabited. More than half of the
total number of burrows in the Lena-Khatanga territory (6000 bur-
rows) are located in the northern subzone of the tundra; the shrub
tundra is the most poorly inhabited. There are many burrows in the
Lena river delta. Now and then, burrows are met with at a distance
of 200—250 m to 0.5—1 km from each other, but are usually scattered
tens of kilometers from each other (A. Romanov, 1941). In unfavor-
able years, the number of arctic foxes here in winter is not large. In
Anabarsk and Ust’-Yansk regions, an accumulation of burrows is
noticed only in some places. In the upper Baranikhi river, near Kolyma,
F. Vrangel’ (1841) found numerous burrows of arctic foxes in the
past century.

In the years of low numerical level in the Lena delta, the baited
traps with an effective radius of about 300 km? took in from 4—6 to
30—40 arctic foxes in the winter of 1932 (Freiberg, 1939); the prob-
able population density—up to 1.3 polar foxes per 1000 ha. When the
total number of arctic foxes was high in northern Yakutiya in the
lower Indigirka and Kolyma in the winter of 1946/47, the average
hunter in one day’s harvest encountered up to 13 arctic foxes; the
ordinary hunter caught about 50, and the best one caught 183, for the
season. With low numbers in the 1948/49 season on a journey of 20—
50 km, both in the northwestern and northeastern regions of Yakutiya,

298

hunters rarely met with more than 1-4 arctic foxes; of them the average
caught 10 for the winter, and the best 20-30 animals. Arctic foxes are
quite numerous on Wrangel’ Island (Mineev, 1935; Sergeev, 1936).

In extreme northeastern Siberia (Magadan district), the average
density of fox populations in tundra subzones differ by a factor of 10-
fold (Table 20) (Shustov and Belozorov, 1959). In Bilibinsk region of
this district in the winter of 1948/1949, in a year of quite large
numbers of animals, the best hunters took up to 40 animals (Chirkova,
1955). The arctic fox was scarce in the 30’s on the Bering Sea coast
from Korf Gulf to the mouth of the Apuka river (Koryaksk Nats.
Region), it was hunted singly and rarely in tens in each region
(Samorodov, 1939).

Estimates of arctic foxes on the Commander islands are known
from the first decades of the present century (Tables 20 and 21). On
Bering Island, the total number of arctic foxes changed through the
years from approximately 2 to 4 thousand, and on Mednyi Island was
in the range of one thousand; the population density on the latter
island was 2 or more times greater. The considerably great density of
the arctic fox population on the Commander islands, as compared to
the mainland, depended at the end of the past century and the begin-
ning of the current one, on the abundance of marine animal carcasses
and from the 20’s—on the systematic provisioning of the arctic foxes.

For the historical period in the USSR*, the number of arctic foxes
changed to the reduced side, chiefly on isolated islands where there
were numerous blue foxes (Krasheninnikov, 1755; Steller, 1774, 1781;
Gondatti, 1898; Nosilov, 1909; Formozov, 1929). Mention by
Krasheninnikov concerning the large quantity of arctic foxes on
Kamchatka, and Steller’s description (1774) of the innumerable herds
of these animals in the middle of the 18th century, are known; the
latter killed in one day up to 70 arctic foxes with an axe. The large
number of arctic foxes on the shores of the Bering Sea in the past
century and earlier, when arctic foxes in “hundreds ran along the
shore” depended upon the abundance of marine animals, on the car-
casses of which, the arctic foxes were feeding.

There were large numbers of arctic foxes on Novaya Zemlya at
the end of the 80’s and on Kil’din Island in the 70’s of the past
century with the first permanent appearance of humans. One needs to
assume that the number of arctic foxes on the Kola peninsula and in

*From the citation dates, this apparently refers instead to Russia—Sci. Ed.

213

299

the northern Far East, also greatly decreased, where the best coastal
places for polar fox settlement are now densely inhabited by man.
There are some data about significant decreases in stocks of arctic
foxes in Sweden (Lost, 1955).

Habitat. Arctic foxes prefer open tundra areas where it also re-
produces. Only in the years of high numbers and insufficient
unoccupied habitations do arctic foxes breed among shrubby vegeta-
tion. The southern border of the denning area coincides with the
northern limits of distribution of tundra-edge forests. Southward,
burrows of arctic fox are only found in broad expanses of montane
tundra. In connection with the climatic warming in the Arctic in the
course of the last series of decades, a shifting of arctic foxes from the
southern belt of tundra to more northerly subzones is observed
(Skrobov, 1960, 1960a).

The main factors which regulate the distribution of the species in
the period of sedentary life are the relief of the locality and the food
conditions. The most suitable places for denning are hilly landscapes
of dry open watersheds and sources of пуще! and streams; smail

Fig. 83. Tundra along the Khatanga at 72° N. Lat., Taimyr. Core fox habitat in
summer. Photograph by A.F. Chirkova.

214

300

sandy knolls in open tundra; the upper parts of high terrace slopes of
creek valleys, the borders of terraces above floodplains, elevated lake
shores, islands and seashores. Near the southern boundary of the
range in the summer time, arctic foxes den in treeless rocky slopes
of mountains, mainly northern exposures, and also in high open water-
sheds. Burrows here are always situated on elevated sections of tundra.

Places of denning and feeding usually do not coincide. Denning
regions change annually. In years with an abundant “yield” of lem-
mings, a large number of arctic foxes live in the arctic and subarctic
tundras. With a depression in rodent numbers, arctic foxes remain on
the seashore, in the southern subzone of tundra and forest-tundra; in
optimal years, the arctic fox usually does not live in the latter.

Near the seashore, in connection with better security of food, the
arctic fox population is more permanent than in other zones. As is
known, habitats are abruptly changed seasonally throughout the year
in connection with migrations. Places where small rodents are found
in superabundance serve as feeding biotopes; at any time of year in
summer, moreover, such places are those where waterfowl occur. In
winter, arctic foxes also feed along the shores and in areas where
willow ptarmigan [Lagopus lagopus] are concentrated.

In the Kola peninsula, breeding places of the arctic fox are re-
stricted to hilly landscapes at a distance of 40—50 km from the sea.
The arctic fox constructs the burrow in bearberry-reindeer moss,
hilly, lacustrine tundra and on alluvial meadows, and forages chiefly
in willow-grass, sedge, lacustrine, and rocky stand-types, and along
the seashore. On Kil’din island, it often lives in coastal biotopes
(Formozov, 1927; Dubrovskii, 1939; Zolotov, 1940). On Novaya
Zemlya, arctic foxes densely inhabit the low-lying Gusinaya Zemlya
with its hilly landscape in the southwestern parts of the island; fewer
arctic foxes occur in montane tundra at Matochkin Shar; in the polar
part of Severnyi Island which is covered with an ice shield, arctic
foxes are found only along the sea coast (Zhitkov and Buturlin, 1901;
Nosilov, 1909).

The dispersion of burrows in Kaninsk, Timansk and Malozemel’sk
tundras is closely connected with sandy-hilly tundra (Shibanov, 1951;
Maksimov, 1959). In Malozemel’sk and Bol’shezemel’sk tundras,
93% of the burrows are located on hillocks, knolls, slopes of ravines
and along the shores of water bodies. Of the burrows 75.5% were
placed among types of tundra vegetation, of which 22.5% were in
moss, 18% in lichen, and 16.1% in shrub formations. 13.8% of the

301

burrows were located within marshy tundra, of which the most fre-
quent type—9.3%—were found in the vicinity of marshes in
grassy-tundra, 10.5% were found in shrub tundra, and in meadow
associations, 0.2% of the burrows (Skrobov, 1958).

On the Taimyr peninsula, the number of burrows dies away to-
wards the north, as a function of the transition from the tundra zone
to the arctic desert zone. On the Chelyuskin peninsula, burrows are
absent. Burrows are rarely found along the coast of the Kara Sea, are
relatively frequent along the northern shore of the peninsula and
along the Nizhnaya Taimyr river. Along the entire western shore of
the peninsula, the coastal and pre-delta sections, and river and creek
banks are the areas most densely inhabited by arctic foxes. On the
eastern coast and on Begicheva Island, their burrows are often situ-
ated in hilly tundra. In the north of the peninsula, the arctic fox
reproduces among cotton grass-sedge-moss tundra, where there is a
rich fauna of rodents and birds, and also along the edge of stony
deserts, along ravines and in glacial cirques on the montane plateau
of Byrrang. Burrows are dispersed among dryas patches and grass-
moss in the arctic semidesert (Koshkin, 1937; L. Lopov, 1939; Shastin,
1939; Sdobnikov, 1959).

In the Lena-Khatanga territory, satisfaction of the basic nutrition
of the arctic foxes is associated with tundra intersected by a dense
network of small creeks, lakes and /aidas*, lake depressions, and the
seashore, which are the main feeding lands of arctic fox (A. Romanov,
1941). On Wrangel’ island, the arctic fox lives among sandy hillocks
and feeds along the seashore.

In the basin of the Anadyr’ river, the arctic fox thrives abun-
dantly in the continental and maritime arctic level tundra, but are also
abundant in the hilly shrub tundra. There are small numbers of arctic
fox in montane lichen-shrub tundra and in the forest tundra, and a
negligible number in the level shrub tundra. In alpine tundra the
animal is absent. In the montane tundra region of the southeastern
part of the Koryaksk National Region, the small number of arctic fox
here are restricted to tundra, the seashore and among shrubs in river
valleys (Mineev, 1935; Mikhel’, 1937; Gassovskii, 1939; Samorodov,
1939).

In Magadan district, the density of foxes is highest in the north-
em maritime subzone; in the southern maritime subzone, its density

*Openings in the forest-tundra landscape—Gen. Ed.

215

216

302

is less, and in montane sections of the central interior subzone which
are far from the sea, its density is lowest. In 1000 ha of different
types of hunting lands the distribution of arctic foxes is, on the av-
erage, as follows. In flooded shrubby and meadow tundra—biotopes
favorable for reproduction—S arctic foxes. In complex marshes, in
the lower courses of large rivers, and along level watersheds and
maritime plains with abundant lakes—the main biotopes of arctic fox
in the period of reproduction and raising of young—2.25 arctic foxes.
In the low-lying and transitional marshes and marginal shrub tun-
dras—1.3 arctic foxes. In hummocky, cotton grass-sedge, and oiser
shrub tundra of the northern subzone—0.75 arctic foxes. In tundra,
shrubland and marshes of the southern and central subzones of the
arctic tundra—0.5—0.25 arctic foxes (Shustov and Belozorov, 1959).

In Kamchatka, the arctic fox inhabits the zone of sparse stone-
birch forests. On the Commander islands, it dens in maritime hillocks
and rocks and, in fewer numbers, lives in the interior of the islands.
It feeds, mainly, in the Jaidas, cliffs of bird colonies, and along the
banks of rivers; in autumn, in tundra and on hills. Throughout the
year, they frequent human settlements (Suvorov, 1912; Barabash-
Nikiforov, 1937).

In winter, the arctic fox is often encountered on ice. During the
time of migration to the forest zone, they stick to open places—triver
valleys, watersheds, fields and marshes.

Food. The arctic fox, like the red fox, is an omnivorous beast, but
more voracious and less selective; it is poorly provided with food and
is frequently hungry for long periods. Within the USSR, more than
125 species of animals and 25 plant species are known to be eaten
by the arctic fox.

The nutrition of the arctic fox in continental tundra and on the
seashore differs sharply. Already investigators of the past century had
emphasized the dependence of arctic fox on lemmings (Pakhtusov,
1842; Krivoshapkin, 1865; Middendorf, 1869; Tret’yakov, 1871;
Silant’ev, 1898). At the present time, numerous investigations have
shown that the main food of tundra arctic foxes are small rodents,
which constitute in some seasons and years more than 99.0% of the
stomach contents and feces; mainly the species represented are Nor-
way, Ob and hoofed [= collared] lemmings (Lemmus lemmus, L.
sibiricus, Dicrostonyx torquatus). Gray (genus Microtus) and red
[backed] (genus Clethrionomys) voles are eaten by arctic fox in smaller
amounts; these rodents are met with in the arctic fox’s diet mainly in

215

303

the southern tundra belt, in forest tundra and in the taiga zone (during
migration). In places of water vole (Arvicola terrestris) abundance,
arctic fox feeds also on them; they rarely catch arctic hares which are
particularly characteristic of the shrubby growths on the
Bol’shezemel’sk tundra (up to 43% in studies).

Birds are caught more often in spring and summer (in Yamal up
to 52% occurrence in analysis). They destroy nests, eating the eggs
and nestlings; in the second half of summer, they hunt molting geese,
most frequently the white-fronted goose, and rarely, ducks. In autumn
and winter, they catch willow and rock ptarmigan [L. lagopus, L.
mutus], rarely snow buntings [Plectrophenax nivalis] and longspurs
[Calcarius lapponicus], and still more rarely, birds of prey wintering
in the tundra. Quite often they eat fish (up to 41% occurrence in the
food analysis), usually refuse of fish harvest or bait; it sometimes
catches migrating salmon during spawning time,

Fig. 84. The shore of Kara Bay (Kara Sea) at the beginning of winter. A place of
massive migration and harvest of the arctic fox. Photograph by A.F. Shirkova.

304

Wild and domestic reindeer are of great importance in the nutri-
tion of tundra arctic foxes, herds of which arctic foxes usually follow.
In the diggings [feeding craters] of reindeers, the arctic fox catches
small rodents; near the reindeer herd, ptarmigan are concentrated;
fallen reindeer become food for arctic fox (up to 68% in food remains
in famine years). In winter, arctic foxes eat the feces of reindeer.
Sometimes, arctic foxes pursue reindeer weakened by a prolonged
period of ice-covered ground or injured reindeer; they devour lumps
of snow moistened with the blood of these animals; they lie around
the dying animals in expectation of their death. Cases are known
when arctic foxes killed and ate new-born reindeer fawns, even in the
presence of their mothers.

Most of the above-listed foods are eaten by the tundra arctic fox
only in the absence of lemmings. One must consider them secondary.

Invertebrates, especially marine, are insufficiently known. In-
sects occur rarely and in negligible quantities in the food of the arctic
fox—heetles, bumblebees, wasps, grasshoppers. Extremely rarely, the
arctic fox eats amphibians (grass frog on the Kanin peninsula). Weasel,
ermine, shrews, insects and amphibians must be assigned to the class
of accidental and emergency food, eaten in the years of lemming
scarcity.

Cannibalism is developed to a strong degree among arctic foxes.
In some years, more than 50% of those animals falling into traps,
were found to be chewed. Cases of eating dogs and red foxes are
extremely rare!’.

Arctic foxes inhabiting islands and coasts feed on the flotsam of
the sea—carcasses of marine animals and birds. They gnaw a way
through the whole system of a whale’s carcass and sometime do not
come out of it for some days (Nosilov, 1909; Mineev, 1935): they
gorge themselves with blubber until they vomit. On the littoral, they
gather cast-up cod, haddock, sea-hen, capelin, arctic cod, navaga
[Eleginus navaga], smelt and other fishes. There, they also gorge on
echinoderms, crustaceans, molluscs, sea-cucumbers, ascidians, sponges,
hydroids and others. In a period of massive consumption of sea ur-
chins, the teeth of arctic foxes on Mednyi Island are stained violet in
color. Bird colonies are preferred foraging places for arctic fox. They
capture black guillemots [Cepphus grylle], murres [Uria sp.], fulmars
[Fulmarus glacialis], razorbills [Alca torda], gulls and even the
stronger skuas [Stercorarius], cormorants, goosanders [Mergus] and

170.4—2.0% in analyses of food (Pereleshin, 1943; Skrobov, 1958).

РИ

218

305

others. In one feeding, a hungry arctic fox may eat up to 0.5 kg meat,
fat and fish (Skrobov, 1958). The gastro-intestinal tract of the arctic
fox fully rids itself of food remains within 24 hours (Pozgnyakov,
1957).

Arctic foxes feed also on plant food; in the tundra on various
fruits—crowberry, blueberry, whortleberry, bearberry, cloudberry,
various species of sedges and grasses, and on the seashore, on
seaweeds; the significance of the latter is not entirely clear, since it
is evident that they are not digested at all. Arctic foxes chew on
russula mushrooms. In the alimentary tracts of 898 arctic foxes were
found 78 different species and forms of plants taken during feeding
on other food materials (Pereleshin, 1943). For the arctic fox, as for
the red fox, ingesting inedible material—discarded reindeer skin, raw
strips, rags, string, paper, woodchips, crushed pieces of coal, pieces
of soap, as well as pebbles, sand, glass fragments and others—is
characteristic.

On moving in autumn and winter to the ice of the Arctic Ocean,
arctic foxes (especially the old ones, for which independent foraging
is difficult) usually follow polar bears, gathering its food leavings,
and also remains of the food of ringed seals; they gnaw at the eruc-
tations of the walrus—a whole pile of indigestible food. Arctic foxes
return in spring from the ice seriously emaciated.

The prevalence in the diet of these or other food items depends
upon the season of the year. Tundra arctic foxes most frequently
consume lemmings at the end of summer, in autumn and in the first
half of winter, i.e. at the time of their mass appearance. The most
stressful time of the year in tundra is the end of the winter and
beginning of spring. The long duration of ice-cover, often in the
second half of winter, results in the emaciation and death of reindeer,
thus providing additional food for arctic foxes; at that time, the ice
crust which forms renders difficult the excavations of voles and causes
the death of small rodents. At that time, arctic foxes with broken
claws are not infrequently encountered. In spring and summertime,
the possibility of the foxes’ feeding at the expense of birds increases.
In the second half of summer, the molting birds appear, and in au-
tumn berries ripen.

The diet of the arctic fox varies in different geographic regions.
In Iokan’go-Ponoisk tundra of the Kola Peninsula (Table 22), at the
time of rodent reproduction, the arctic fox most frequently feeds on
Norway lemming (to 96% of all data), they often eat gray and red-

306

Time of year

Types of food

September
November

Be
$

Rodents

Carcasses and remains of reindeer

Willow ptarmigan (Lagopus lagopus)

1 Guils and eider [ducks }

Seals

Sea urchins
Starfishes
Sea cabbage
Crabs

НЕ

nae
LR RE ee aT aC ОЕ

i>
те [

217

и.
Be
a

Е oe cn yan ntl
Beta setae AN an Oe
Ea i a BZ
Е Doai Ci a nvee 525255280

Fig. 85. Change i in the composition of arctic fox diet during the course of the year.
A—amainland arctic fox of Kola peninsula (Zolotov, 1940); B—island arctic fox
of Kil’din Island, Murman (Formozov, 1929; Zolotov, 1940).

backed voles, birds and insects. On Kil’din Island in the summer of

1927, foxes mainly existed at the expense of the shore fauna due to
the scarcity of rodents (Fig. 85; Formozov, 1929).

In the northern parts of the Kanin Peninsula (Tables 23 and 24),
feces collected near the den burrows of arctic foxes contained, almost
exclusively, mammalian remains. The majority of them belonged to
collared and Siberian lemmings, and in lesser quantity, voles and
others. In a year when lemmings were absent, cloudberry was pre-
dominant among plant foods. In Bol’shezemel’sk and Malozemel’sk
tundras (Table 23), the diet of the arctic fox consisted most frequently
of mouse-like rodents, including Siberian lemming, and when the
latter was in small numbers, narrow-skulled vole (Microtus gregalis)
and water vole (at the mouth of the Pechora); moreover, white hare
and, among birds, chiefly willow ptarmigan (probably the latter, as
well as hares, represented bait from traps*). Remains of shrews

*The Russian phrase, “nazhivky iz samolovov”, probably refers to bait from a
dead-fall trap; see Figs. 96, 97—Sci. Ed.

307

(winter, 1957/58, up to 8% occurrences) were found in arctic foxes
caught near Karsk Bay. Fox stomachs in the winter of 1956/57, when
lemmings were absent, contained much garbage and household refuse.
Around burrows of the arctic fox in summer are often found wings
and bones of willow ptarmigan, and rarely, of ducks, small shore-
birds and small passerine birds and among mammals, the white hare
(Skrobov, 1958).

The nutritional condition of arctic foxes in northeastern
Bol’shezemel’sk tundra depended on the nutritional conditions of the
year, as well as on sex, age, and physiological condition. In winter
1956/57, distinguished by low numbers of small rodents and large
numbers of arctic foxes, males were in better condition in autumn,
while during winter they lost weight. Females, especially those with

Table 22. Diet of arctic fox on Kola Peninsula (in % of occurrences in number
of investigated objects)

Place of collection, Iokan’ go-Ponoisk tundra Kil’ din

season, year Pao Pomp lols Fa 4 ue Island,
AE SRS Se Gea a 1927,
Food Зиттег* Winter'® August’? Summer’?

Small rodents 80 20

Lemmings 96 92

Gray voles [Microtus] 46 15

Common and Northern red-backed 42 13

vole [Clethrionomys]

Arctic hares [Lepus timidus] 1.3

Shrews [Sorex] ТЗ

Carrion (reindeer) 9 18

Birds 42 29 16 40

Fish 10

Insects 29 Less than 5

Marine animals of the littoral zone 2522

Crustaceans 5)

Molluscs я

Plants 7 4 Less than 5

Refuse 11

Number of small rodents Very Low
low

Г.М. Borodin.
'Zolotov, 1940.
°Formozov, 1929.
*|Berries.

*2Mostly sea urchins.

9$
8 С
ov
ОТ
VL
[УС

LS/9S6l
JOWUL М

SBUIPULJ JO %

SOUIJSOJUI PUB SYORUWIO}S 06

че АЯ 151еУ

10 T 67
O¢

928° 920° L
Tl 90
LG SL
€s Cer
VTL ОТ
89/1961 95/6661

JOVI AA

JYSIOM JO %

SYOBUIO]S CS]

petdpun} 1$.[9\1э20 |8
pue ys jawezays jog

£0
СИ СФ
го
СИ v9
214! |8эи
VCL €'€8
Lv6l 961
Jowwins
$8ириц JO %
SJUSWIIIOXE +09 755

севЗитиэа uluey
019159 4310 №

(сшешол JO 3481эм [6303 JO % UI рив
рэзе8 элит ит S9IUAIINIDO JO JOQUINU [8303 JY) JO % Ul) JIAISIP YS Дэ8иецхлу илэцзлои JO влрипз и! JIIP хо} эволу “ET GEL

$и2998151.1)
$12954]

ysl

$889 $. рия

риа

зтецние suey
(лээритэл) ио1ле
SMOIYUS

ore SHUM
SJUIPO] еше

мени

pood

18 ‘uoseas
‘sjyoofqo jo зипоше
‘UOI}D9][0D JO э284

syalqo

‘URSIUIE}d MO[[IM JO SUIBWOI A]}SOP[gz
‘6561 “TE 32 BAOYNYOcz

"8061 “AOQOLS yz

8561 “AOURQIYS,,

Joquinu ul
эзволоэр dievys ‘/p6| ul

MOT] wnipoy MOT шатрэш ‘961 Ul ззиэрол Пе JO 1эдщям
001 001 001 001 001 % Ul [830.1
5'9 10 э$пуэл pjoyssnoy
LEAS 10 91 ($12430 pue spunodwod [ejouru

‘sjou ‘Ioyyeo] ‘sde1) $1ээ(4о э1атрэи]
LY (as sjUe|d
8 rl €0 19:8 |3эи 2198 18эи хо} эцэлу

70 10 пои

219

220

310

litters, were more emaciated in autumn, but in winter, they fattened
up; only a few specimens were characterized by severe emaciation—
purplish color of muscles and great projection of bones. At the same
time, arctic foxes which were, apparently, feeding on the coast, were
noted to have large fat deposits—subcutaneous fat on the sides up to
2—3 cm, and in the form of layers in the muscles. From one male, up
to | kg of fat was removed, which represented 1/4 of its carcass
weight. In the following winter 1957/58, distinguished by abundance
of lemmings and low number of arctic foxes, the young females were
very well nourished, but those which reproduced became of lower
than average nutritive condition (Chirkova et al., 1959).

Foods of the arctic fox on the islands of Novaya Zemlya are
varied. The animals move back and forth everyday from the tundra
in which they catch lemmings, to the coast where they feed on marine
flotsam. In rivers, they catch chars and live around the carcasses of
marine animals for long periods in snow burrows. Hunting in bird
colonies, they destroy completely the accessible nests. In Franz Josef
Land, the arctic fox feeds mainly at the expense of bird colonies
(Nosilov, 1909; Esipov, 1935).

Yamal arctic foxes, far from the coast (Table 25), fed exclusively
on terrestrial mammals (in 1939—1942 up to 99%—100% of occur-
rences in diet; Dunaeva and Osmolovskaya, 1948), and sometimes
uninterruptedly on mouse-like rodents. In summer 1933, in the Tambei
river basin, the latter formed 99.6% of occurrences in feces at the
burrows (Tsetsevinskii, 1940). They were mainly Siberian lemming
(up to 79.5% of occurrences from total sample amount and up to
92.5% of all rodents in northern Yamal), and fewer collared lem-
mings, and also gray and red-backed voles. The importance of
lemmings, especially the Siberian, increases as one moves northward
on the peninsula. At places in the floodplain of the Ob’ river and Ob’
Bay, water voles occupy some place in the diet of the arctic fox. The
portion of birds in the diet of tundra foxes with lemmings abundant
is not great even in summer, but in individual years in the taiga belt
of the Yamala-Nenetsk National Region, it increases greatly (to 50%).
Here, the arctic fox gorges on fish, mainly the sorts of refuse from
places of fishing harvest. In a year of abundant berry crop and few
lemmings, plant food is of great importance and also carrion.

Great fluctuations in the dietary composition is observed in the
tundra arctic fox, depending on annual abundance of lemmings (for
example, winter 1932/33 and 1933/34—Table 25: Tsetsevinskii, 1940).

222

223

311

In winter 1938/39, in southern Yamal (Korzinkina, 1946), the occur-
rence of rodent remains in the arctic fox diet was observed to grow
monthly in connection with subnival lemming reproduction. Bird re-
mains also declined. With an abundance of rodents in autumn 1939
and their gradual disappearance in winter 1939/40, the occurrence of
rodent remains in the stomach contents diminished monthly; at the
same time, the arctic fox fed more often on reindeer carrion and
caught birds.

On the Taimyr peninsula, the principal food of the arctic fox also
consists of small rodents. In August 1931 on the right bank of the
lower Khatanga river, among the bones of mouse-like rodents col-
lected near an old den burrow, 85% of the skull remains consisted of
collared lemming, and 15%—Siberian (А.Е. Chirkova). In winter
1932/33 in the neighborhood of Khatanga village and a series of
eastern stations* (Table 26; Skalon, 1936) with a lemming die-off,
small rodents occurred in 68% of the dissected stomachs of arctic
foxes; predominantly voles—M. middendorffi and М. hyperboreus
and the Siberian lemming. The remains of reindeer carrion, fish mainly
from baits, and also willow ptarmigan and some other birds consti-
tuted the three other types of food; each of them was found in nearly
1/3 of the stomachs. The considerable mixture of food and inedible
human household refuse was also an indication of the lack of small
rodents in that winter.

On the southwestern coast of Taimyr and on Begichev island, the
principal food of the arctic fox was also recognized as lemmings,
especially the Siberian. On the Chelyuskin peninsula, in a lemming
year, the remains of collared lemmings—the only one occurring here—
constituted the principal content of tundra arctic fox feces. In years
when rodents disappear, the diet of the arctic fox mainly consisted of
marine flotsam which is not present here each year owing to the
different times of ice formation, and of the leavings of the polar bear.
Arctic foxes fed in autumn 1932 on marine flotsam. The foxes also
dug this food out from under the snow in winter, living in a series of
snow burrows. In winter time, arctic foxes here frequently do not get
enough to eat, and in connection with this cannibalism develops among
them. In the famine winter of 1944/45, arctic foxes dug up the lem-
ming carcasses stored from the preceding year, when these rodents

*In the original Russian text, this word is stankoy (gen. pl.; machine-tool);
apparently a misprint for stantsov—Sci. Ed.

220

224

312

Table 24. Mammalian species in food remains of the arctic fox on Kanin
Peninsula in summer at burrows (in % of total number of occurrences
in feces; Shibanov, 1951)

Quantity 554 604

Food Year 1946 1947
Collared lemming [Dicrostonyx torquatus] 56.8 48.9
Siberian lemming [Lemmus sibiricus] 21.1 8.4
Wood lemming [Myopus schisticeps] 0.5 0.2
Lemmings, not accurately identified 0.4 —
Root vole [Microtus oeconomus] 3.3 4.0
Northern red-backed vole [Clethrionomys rutilus] -—- 0.3
Voles, not accurately identified 10.4 30.4
Mouse-like rodents, not accurately determined 551 6.6
Shrews [Sorex] — 0.2
Arctic fox 1.8 1.0

100.0 100.0

were abundant, and died after that (Kirpichnikov, 1937; Koshkin,
1937; Rutilevskii, 1939; Sdobnikov, 1958).

In Lena-Khatanga territory (Table 26) during a sharp reduction
in numbers of mouse-like rodents, mainly collared and Siberian lem-
mings were found in the stomachs of half the arctic foxes in winter
1926/27. The foxes took birds and fish more often from lures (A.
Romanov, 1941). In the lower Lena, the arctic fox also preys on
northern pika—Ochotona hyperborea (Kapitonov, 1961).

In the eastern regions of northern Yakutiya (Table 26), the years
in which the investigation of arctic fox diet fell apparently coincided
with few rodents (Krasovskii, 1939). In the delta of Lena, the arctic
fox frequently fed on fish and birds.

The existence of arctic foxes on the Commander islands is asso-
ciated completely with the seashore. Their diet is composed (Table
27) of the carcasses of marine mammals, especially fur seals
[Callorhinus ursinus] (up to 56% in the analysis), birds, echinoderms,
mostly sea urchins, and other littoral invertebrates, fresh fish, and
dried salmon from foodstores, crustaceans, and molluscs. Sometimes,
arctic foxes capture young fur seals from their mothers, or drive away
ringed seals from their breathing holes into the ice and then attack
them. Cannibalism is not observed on the Commander islands. Among
plant food, the arctic fox eats berries in the tundra, laminaria kelp,

(‘рлио2)

zexPOOJ
jejuoweddns

90 *“MO]Aq 99S зтеипие эиыеи|
50 58—81 7—1 1°89 eS Jaapulal JO UOLLIeD
ф 0} dn Gil [хэ4о$] SMO1YS
€0 [sijpaiu 612151] 1э5еэ М
9 0} dn [snpiu зпаэ7! элеч sy М
81-7 рэуциэр! JOU ‘ззиэрол [еб
OGY, рэцциэр! JOU *So}OA
Г 0} dn [5141594491 DJODIAAP] JOA 1938 М
0—9`0 [19$ sdmouolsyjaj)] э1о^ payoeq-poy

Эдо т
9°8 09-F'l pue [1 0} dn 9 € we V [9$ snjous1py] aoa 40
Bee 97
= vISI— С 99] wet [snjonbaoj х4ио150421А | Зчилияэ1 рэле[о)
0'55 ОЭ 9 reO SL veS CO [5712141915 зпшшат] Зитилилэ| UBLIOQIS
8'98 Гб Вии]
7 56 L6— 6L $19 6'76 зиэрол |185

oC V6 JOUUINS 65/8561
"зеииеш 0761-6861 —/ 56/9661 ve/ccol €t/Ct6l 29/5561
JO Sada} JOJULM, JO SIOIUIAA ТЭЛА JOU TOJULM
Pre JO % Ul ‘$48015 086 (0}—WOIJ) SIBIA "599; рие ‘$999; рие $498 101$ $6
еше э|ррищ UI $Эи95 ЭТ SYSBWOJS EQ] SYOBUIOIS 86 ‘рие[$ |Аэяа
ТИР ОТВ О ое
868 ‘веварипз gzlOqUl М
0} 8518} шо44 ‘лэлы 1эдще | ‘влрипз эпоху 1eak ‘uoseas ‘ззоэ[ао Jo
a aaa eS come ae ee Jaquinu ‘U01999][09 JO 99k |
ewe x ewe x

(s}dafqo рэзе8и5элиг JO лэашпи 0} SddU911NIIO % Ul) PULIS] 1А|эа PUL VINSUIUIg [VWLA UO JIIP хо} эполу “SZ 911 ГСС

‘Pa 125

—59эхо} JO иоцециээиоо [е20] 9BeINODUS 0} рипол8 dy} UO рэ1э1е5$ poo} [виэшэ|Ч4п$ 01 siajor Апиэлеа4е „вуишохро4,, рлом uvissny su Ly
"SJUIPOL JOre

‘%IL—VE/ELOL Ul PUL %YOZ—EE/ZEG1 Ul [OOM хо} 94948 “JOAOIIO|g¢

‘yey [RWIUR эцыеш JoyJO pue [Dpidsiy 02044] [еэ$ PIBUIY 2,

‘uesiIUie}d моим K]ISOJ i>

"SpE ‘VAPYSAO[OWSO PUL LAILUNC ge,

EVO] `914 59191 dz

`ОУ6Т “PYSUIAISIOS L gz

‘SE61 ‘ИИА ие

2198 18эи yeas
129148 Алэл suisueyo Зи MO] $зиэрол [[VwWs JO JOqUINN
IZ 6:1 э5пуэл pjoyassnoy
5—0 OI-Z УС (‘232 ‘зрелэцим ‘5зэи ‘1э43е3| ‘$8ел) э1атрэц]
\(Gmeks S6— IL 811 ge ива
OI-L0 595 ГР y'9C хо} эцолу
$19430
рие $883 SIOYIVIJ риа 6'19 sjleq ‘sain, ‘pooy [езиэщэ[94п$
Gl SOSNT[OW
£0 €-$0 Coz C0 Gl 8 Ч
(5—9)
NS 9-18 5—6 (91) (175) 8 (з1эцзеэ} Ато) зрия

(р/ио2 сс 21981)

(‘piuoa)

С Cll хо} O1DIY
sp1iq -Ysy}

‘лээритэл APsop] 542410 pue Ч sjleq ‘Soin, ‘spooj [е}иэщ1э1491$
L0 ГЕ 61 spodoiyiy
СУ $95 95 eet 58 4st
50 ба у 5889 $.рия
vl Е ysl ГЕ С 87 ри
ИС S[eUNIUe JUICY
ВЕ 50 EL ol OVE тээритэл JO UOLLILY)
4 [х240$] $м924$
el [приии! snda7] эле эичм
9:0 [5$4шоио14ц12]5] зэ1ол рэуэе9-рэч
9'0 6:05 [5710421 sajoa Kelp
L'€€ 97 [snjpnbsoy хдио150421а] зицииэт рэле|оЭ
617 С 61 [5721.19 15 зпшшаэт] Зипии| UeLIaqIS
OLE Bet 20 ce sume]
89 ззцэрол [еи$
БЕ ВЕНЕ ee ee ee

9 ¢ Vv € (6 |

УР/СУбТ 1931 6€-LE6I ool Z/9T61 JOUIM че ЕЕ/СЕбТ JOIUIM

"SYSBWIOIS 685 "SYOBWIO}IS /05 “SYORUO}S 091 "34281035
‘uoIdal ‘spurysi $428и10}$ 6651 ‘$99иэ4119э0 ul рипоу poo
YSWA[OY ЭР ysaoyyer “эр еиэ7] Це JO [2303 ‘ejnsuluod лАцие |,
0} ‘Алоэ 3 y
55994924190 [| JO [8101 03 езиеецу ea ‘105895 ‘519[40 Jo

JaquINU ‘UOI}99]]09 JO ээва

еАцихед илэ35$еэЧл0м -виэ7]

(зээиэллиээо || JO
18303 03 $32э[40 рэзе8ц5эли! JO лэашии 0} 2DUAIINIDO % Ul) BAINYBA илэЧзлои рие вии IAWIET, UO JIIP XO} эцолу ‘95 Эа 777

"S[eWIUe эныеш JO зе»

‘sjue|d YSsol4o,

3129 A[ISOWe¢

‘uBSIWIEId моим A]ISOW\g¢

"€S61 ‘ЧО

Т7бТ “AOUBWOY ‘У

"9561 ‘40215
a er онЗЙЗЕНАЙИЙ

[Jews “1ojuIM ит

Joquinu dy} Ul ‘osie] ‘UWNNe

э$вэлоэр dieys (1013 811816 $зиэрол |[е1$ JO JOqUINN
Е А
001 001 001 001 % [BIOL
ЕЕ Е О as Sie eet ees Bae
vty 8°Or 19 CL $428103$ Азия
Or L'8 50 esnjol pjoyssnoy
СУ 67 €9 6 orS 07 sued
ЕЕ а se ee ee

9 6 ф € с |

((p1u0997 21481)

ЗИ

223 Table 27. Arctic fox diet on Commander islands (in % occurrence to number
of investigated objects to total of all occurrences)

Place of collec-
tion, number о —

objects, season, 194 534 250 296
year stomachs, stomachs, to stomachs, to stomachs, to

Bering Island Mednyi Island

to the number of total of all number of
total of all investigated occurrences, investigated
occurrences” objects, 1924/2544 objects,
Food 1929/30 1930/3245

Small rodents 1.1 0.4
Marine animals 0.2 56 6.8
Birds р Dep) 4 39.5
Fish 38.4 10.3 14.9
63.542
Littoral marine 32.4 159 20 30.1
organisms
16.2
others
Crustaceans 3.4 253034
Molluscs 258 1.1 | 23.0
Supplemental
foods, lures, baits, Day 8 13.5
household refuse
Plants 10.3 8.0 Des 14.5
19.143
Inedible (rags, leather, 0.6
nets, minerals, etc.)
Empty stomachs 10.5 5.0 8
Total in % 100 100

“Cherskii, 1919—1920.

BLavrov, 1932.

“Freiberg, 1929.

‘SBarabash-Nikiforov, 1937, 1939 (diet of tundra and not bait-fed arctic fox).
“Octopus, dvuustika*

“Dried fish—salmon

“8Sea cabbage.

“Mostly sea urchins.

'Insects—0.7%.

*Lit., trematode fluke; meaning unclear in this context—Sci. Ed.

318

rockweeds [Fucus] and other seaweeds of the littoral (Cherskii, 1919—
1920; Freiberg, 1926; Lavrov, 1932; Barabash-Nikiforov, 1937a,
1939).

There are age and sexual variations in the nutrition of arctic
foxes. Juveniles more often feed on lemmings. Among the Khatanga
arctic foxes (Skalon, 1936), the lemmings are more often eaten by
females than by males. Males caught mainly gray voles and also fed
more on reindeer carrion or on baits of birds and fish and as a whole
they did not show selectiveness in food. East of the Lena (Krasovskii,
1939), lemmings were also mostly found in the food of females. On
Belyi island, differences in the diets of males and females were not
observed.

Home range. A definite home range is characteristic of the arctic
fox only in spring and summer, during the time of reproduction and
growth of the young. On the Commander islands, at the time of den
establishment and then the emergence and raising of the young, the
arctic fox occupies a permanent territory sometimes demarcated be-
tween natural boundaries in the form of streams, knolls and other
particular relief features. In the event that a “foreign” arctic fox
intrudes, it is repelled by the parents; in this case, even the weakest
female can drive away the stronger newcomer. However, the area
does offer complete security to the whole family, as the parents go
for food to the seashore or to other regions (Cherskii, 1919—1920;
Boitsov, 1937; Barabash-Nikiforov, 1939). Thus, the home range is
as much a breeding as it is a feeding territory.

The dimensions of the selected territory vary according to the
locality and abundance of food. In rugged terrain, the areas are smaller
than on the plains. In the delta of Lena, the area inhabited by a family
in the summer is from 5 to 30 km’. In the northern part of the
Kharaulakh range, dispersal of the arctic fox from the burrow in June
1927 did not exceed 5 km; the same was also observed on the plains
in the lower Anabara river in 1934. Both years were characterized by
low numbers of lemmings. On the Taimyr, in the region of Ozhidanie
bay, in the year of a good “yield” of arctic foxes (1947), the area of
their breeding territory comprised not less than 16-25 Кт?: 3 burrows
with litters were found at a distance of 4-5 km from each other (A.
Romanov, 1941; Sdobnikov, 1958). Breeding territory on the Kanin,
defined as the distance between burrows, was equal to 2-14 km’,
sometimes more in North Yamal—up to 25—30 km/, and in the Lena-
Khatanga territory—from 20 km? and more. On the Bol’shezemel’sk

225

319

and Malozemel’sk tundras, in 7 centers of arctic fox concentration in
an enormous territory of 16,500 km’, in a comprehensive tally each
2 km? contained only one burrow on the average (Skrobov, 1958).
With food abundance and dependability, litters of these animals are
sometimes placed in immediate proximity to each other, as was ob-
served on the Yamal, Kanin, and in Belyi and Mednyi islands
(Iokhel’son, 1898; Freiberg, 1926, 1939; Tyulin, 1938; Tsetsevinskil,
1940; A. Romanov, 1941; Shibanov, 1951; Sdobnikov, 1958; Skrobov,
1958; N.N. Spitsyn).

As soon as the young emerge on the surface, arctic foxes often
roam from one burrow to another, especially if they are disturbed. On
the Commanders, parents accompany the young to the seashore to
temporary shelters (Cherskii, 1919—1920; Barabash-Nikiforov, 1937а;
Maksimov, 1959).

After the young arctic foxes establish themselves at the end of
summer and beginning of autumn, the boundaries of the littering sites
cease to exist. At this time, the arctic fox commences a nomadic life.
The distribution of arctic foxes after that depends on the abundance
of food in autumn—winter period. In the years with bad supply of
food, the arctic fox moves for some hundreds, thousands and even
more kilometers. In years of lemming abundance in continental tun-
dras, arctic foxes in summer restrict themselves to region of summer
denning during winter also, and their individual ranges at time are not
large. In the Lena delta in a year of lemming mortality, arctic foxes
which remained to winter in the tundra moved for a distance of up
to 10 km in the direction of baited traps (Freiberg, 1939; A. Romanov,
1941).

In the Bering and Solovetski islands, constancy of occupation
from year to year of different places by definite groups of arctic foxes
distinguished by their fur color and behavior was noted.

Burrows and shelters. The choice of a place for construction of
the burrow, and then its use for many years, depend on many factors.
Chief are the dryness of the soil and the absence of proximate subsoil
water and permafrost, slope exposure, the lookout possibilities of the
place, and food supply in summer time. Arctic foxes’ burrows are
usually located on elevated dry interfluves, on small hills, precipices,
etc.

In the mountains or in outcrops of bedrock, the arctic fox digs a
burrow in a fine-grained layer, and if it is absent, they settle in rock
clefts and between plates of montane bedrock. On Kil’din Island,

320

burrows and dens of arctic foxes are concentrated in a coastal belt 1—
2 km in width or narrower; here arctic foxes live at various altitudes,
from some tens of meters above the water to the extreme tops of the
plateau. Burrows, dug in sand, were abandoned because of frequent
human visitation and their destruction, and the foxes burrowed within
rocky areas under plates of bedrock (Formozov, 1929).

The arctic fox also uses the burrows of other animals, for ex-
ample, the marmot [Marmota camtschatica] (Kharaulakh range; V.I.
Kapitonov). In Magadan district, it burrows in places with old ground
squirrel colonies with softer, drier soil (Shustov, Belozorov, 1959). In
seabird colonies on Novaya Zemlya, arctic foxes dig burrows in the
guano (Nosilov, 1909).

In tundras where wind force is great, sheltering the burrow from
it is important. In the northeastern Bol’shezemel’sk tundra, burrows
are more often placed at a height of 2/3 up the slope, not exceeding
12 m above water level of the nearest water body. In such a place,
wind force near the burrows is reduced by 32% of its force on the
slope crest (Danilov, 1958). In the zones of open tundras in the
northern belt, the arctic fox prefers south slopes, without deep snowy
faces, which thaw out earlier in spring, and which are protected from
northerly wind in summer time. In the Bol’shezemel’sk and
Malozemel’sk tundras, of the 483 burrows inspected, 26.5% were
located on the southern side, 17.8% on the eastern, 14.3% on the
southwestern and 11.3% on the northern side. To the north and near
to the coast, the number of burrows on south slopes increases. Thus,
in maritime tundra of the Taimyr peninsula, almost all burrows are
located on slopes of southern exposure, while in the upper reaches of
the Khantaiki, Kureiki and Kotuya rivers below 67°30° and 68° М.
Lat., arctic fox burrows are restricted to the northern slopes (N.
Naumov, 1934; Sdobnikov, 1958; Skrobov, 1958).

Burrows are constructed in the vicinity of water bodies, usually
within 100—500 т. Paths lead to the water. In the southern zone,
burrows are located near growths of shrubs in which young arctic
foxes hide from enemies and bad weather, and where they can also
find food in-the form of ptarmigan and small rodents.

The arctic fox constructs permanent dwellings, usually in sandy
and sandy loam soils, more rarely in sandy-gravel mixed with pebbles
and still more rarely in clay soil. In summer, the soil temperature in
burrows is some degrees higher than outside. However, Middendorf
(1869) encountered in the Taimyr peninsula burrows in which even

321

in the middle of summer, icicles hung like stalactites, with arctic fox
wool frozen to them.

The arctic fox digs burrows of no great depth, from 0.5 to | т,
rarely more. On Belyi Island, in the Taimyr and in the Lena-Khatanga
territory, they are distributed 10—100 cm above the permafrost (Tyulin,
1938; A. Romanov, 1941; Sdobnikov, 1958). In places with burrows,
as compared to neighboring areas, the level of permafrost is lower.
At Cape Osten-Saken on the northwestern shore of the Taimyr, in a
place where burrows are found, the permafrost level is restricted to
a depth of 65 cm (bottom of burrow to 55 cm), while to the side of
the burrow, [frost] depth is 25 cm.

Complexity of the burrow increases through use from year to
year. As a result of quite rapid breakdown of burrows, arctic foxes
build new passages and the center of the lair gradually shifts. The
breakdown proceeds as a result of settling of friable soils through
constant digging; this is also favored by exposure to the air and
trampling of the burrow by reindeer herds attracted thence by succu-
lent vegetation, and also by the digging of humans. The more favorable
place the burrow is located, the more often it is inhabited by a breed-
ing pair. For that, arctic foxes each time undertake a cleaning of the
holes and exit passages. Usually, arctic foxes renew 5—10 passages 2—
8 m in length. In Bol’shezemel’sk and Malozemel’sk tundras, out of
483 burrows inspected, temporary burrows constituted 5.6%; 47.3%
were recently dug and had from 2 to 10 exits; 29% of the burrows
were of medium age and had from 11 to 25 exits and 18.1% had from
26 to 80 exits; these old burrows were, presumably, dug decades ago.
More than 70 exits in one burrow are known in Kanin, up to 60 in
Lena-Khatanga territory, up to 40 in the Commander islands and up
to 30, in the north of the Taimyr peninsula. It happens that two or
more families may occupy one burrow. On the Commander islands,
arctic foxes inhabited one burrow for 15—20 years (Cherskii, 1919—
1920; A. Romanov, 1941; Il’ina, 1949-1950; Sdobnikov, 1958;
Skrobov, 1958; Maksimov, 1959).

In level places, or among low hills, the area of intercommunicat-
ing passages, representing a whole burrow system not rarely occupies
a territory of 100-200, and sometimes 500 m?. On Belyi Island,
burrows were found in the form of systems of communicated pas-
sages with 53 exits occupying 1800 m7’; during the year of this
observation, 11 exits were made. Along the Tiutei river in Yamal,
more than 10 burrows were close to each other. On Belyi Island, in

322

О
®! O02 x3 AC4 855

Fig. 86. Scheme of the construction of an old arctic fox burrow (from A.
Romanov, 1941).
1—primary holes, 2—secondary holes, 3—nest, 4—underground passages,
5—atrine.

an area of 3500 m’, 8 burrows were found. On Wrangel’ island, the
colonies of arctic fox occupy entire sandy hills and are riddled with
countless passages (Mineev, 1935; Tyulin, 1938; N.N. Spitsyn).
227 In old burrow systems, arctic foxes make passages in 2—3 galler-
ies with some chambers for nests. The den in which the nest is placed
is lined with dry grasses and moss. The size of the nest chamber is
from 0.5 to | m in diameter and 30—50 cm in height. The diameter
of the exit opening of burrow ranges from 15—30 cm, rarely more,
reaching in particular cases 80 cm. The cross section of the under-
ground passages is 12—20 cm in height and 15—30 cm in width. As
a rule, the horizontal cross section of the passages and the exits
somewhat exceeds the vertical (Cherskii, 1919—1920; Boitsov, 1937;
Tyulin, 1938; Dubrovskii, 1940; Chirkova, 1940; A. Romanov, 1941).
The area occupied by the burrows, the underground passages and
dens are very dirty because of the remains of food and excrement. As
a result of plentiful manuring of the soil with feces and remains of
food, rich dense dark green vegetation, up to 0.75 m high (in Kola
peninsula) develops. Plant formations on the burrows are highly vari-
able and represent a type of northern herbaceous vegetation. In such
“arctic fox meadow gardens”, tens of higher plant species grow which

323

are absent in the majority of tundra associations. Dandelion, polar
poppy, Alpine forget-me-not, and foxtail grow in Taimyr below the
latitude 75° N. Lat. Near the entrances to burrows mosses (Mnium,
Drepanocladus) grow. Burrows that were occupied by arctic foxes in
previous years are characterized by camomile [Matricaria], meadow
bluegrass [Poa pratensis] and others, or by dense herbaceous growth
mixed with fire-weed and bistort. On abandoned burrows, often only
short fescue [Festuca supina] is found (Dubrovskii, 1937; Zolotov,
1940; Sdobnikov, 1958; Maksimov, 1959). From a distance, it is easy
to determine the location of burrows, which is evident in level tundra
from 0.5 km, and with binoculars, up to 1.5 km. From aircraft, bur-
rows can be seen well from a height of 400—500 п; and from 1000
m, it is possible to map a majority of centers of arctic fox burrows
(Zolotov, 1940; Freiberg, 1940; A.F. Chirkova).

In winter, in periods of continuous snow storms, arctic fox makes
burrows for shelter from driving snow on lee sides. The entrance is
blocked by the blowing snow and the animal remains in the burrow
until clear weather returns. Snow burrows are not longer than 1—1.5
m, widening at the end, of dimensions [width] 25—40 cm and height,
25 cm. The arctic fox also builds temporary burrows near food ac-
cumulations thrown up by the sea (Nosilov, 1909; A. Romanov, 1941
and others).

In years of massive reproduction, when all littering burrows are
occupied, arctic foxes carry the pups outside the burrows—some-
where under the shelter of grasses, where they stay under cover of
grasses, in a pile,of flotsam, or even in the hollow of a large trunk
of the latter (Romanov, 1941).

Daily activity and behavior. The arctic fox is active at any time
of the twenty-four hour period. Scarcity and variability in the food
supply force the animal to wander widely in search of food. In autumn
and in winter, the arctic fox feeds mainly at night; in spring and in
summer, apparently, in connection with the beginning of breeding and
the appearance of their young—during the whole 24 hours. The arctic
fox frequently rests in midday; in clear, windless weather, it settles
down anywhere on a tussock, hillock or slope protected from the
wind, on ice-hummocks or it buries itself in a snow-drift. In spring,
and in the first period after the (re)appearance of the sun, the animal
lies by a dark cliff or on a precipice where it is well warmed. In the
Lena-Khatanga region, a female arctic fox rested in June after eating
from 15:00 to 17:00 during the day and after that from 21:30 in the

228

324

evening; a male, in October arose from rest at about 06:00 in the
morning and retired again at 14:30. On Mednyi Island, in good weather
in the middle of June, a whole family of arctic foxes may be seen near
the burrow early in the morning and again after 17:00 (Cherskui,
1919-1920; A. Romanov, 1941).

While sleeping, the arctic fox rolls itself into a ball, buries the
nose in the base of the tail and covers the belly and limbs with the
tail. Arctic foxes sleep deeply; during periods of noise—wind or ice-
motion—it may be caught by hand (Mikhel’, 1937; Rutilevskii, 1939).
This is easily done during times of hunger, when the arctic fox is
seriously weakened.

The arctic fox is less sensitive to fluctuations in temperature than
red and corsac foxes, as a result of which, at positive [above-freez-
ing] temperature, the general activity of the arctic fox is less than at
negative. By lowering of temperature from +15° to + 5°C, the fre-
quency of breathing of the white fox is increased. In the blue fox,
within the limits of 0-30°C, such dependence is not observed. Gas
exchange and heat production are greater in summer and less in
winter. Daily heat production for 1 kg body weight is 450 kilo-
joules*', in autumn, it is higher and in spring and especially in
summer, it is still higher, reaching 579 kilojoules (Kalabukhov and
Poluzadova, 1946; Kalabukhov, 1950; Pozdnyakov, 1953).

In spring and summer, arctic foxes are most active on the quiet,
cloudy days and in warm weather, with light wind. On cold windy
days, and especially during heavy rain, its activity is sharply reduced,
and the animal conceals itself in a comfortable place. In winter, arctic
foxes do not go out during strong snowstorms, in freezing weather
with severe wind, and on dark nights. On the contrary, they are highly
active on clear moonlit nights, during northern lights and at dawn.
Arctic foxes do not react to light wind, but in a strong one they move
into it. Upon encountering wind, they come to a halt, most commonly
in mild weather. On Belyi Island, they fall in traps usually in calm
weather and during weak and moderate wind speed of 6—7 m/second.
On the Chukotka peninsula, arctic foxes, on the contrary, are more
active and incautious during violent snowstorms and frequent inspec-
tion of the traps at this time yields a large catch (Freiberg, 1934;
Dubrovskii, 1937; Tyulin, 1938; Shundik, 1948; Yanygin, 1948;
Sdobnikov, 1958). On the Solovetsk islands, up to 50% of the arctic

*'According to international system of units (IS).

325

foxes go seeking for food at a right angle to the wind, up to 40% go
with the wind and only 10% go against the wind (Boitsov, 1927). The
same is observed on the Murmansk coast.

As compared to the red foxes, the arctic fox is bold and more
trusting. Steller (1774) mentions how “innumerable packs” of arctic
foxes surrounded a camp of people and became each day “more
courageous and mischievous”. The arctic foxes strewed about not
only provisions which were even stored in barrels but also invaded
people’s living sites, chewed and dragged off clothing, shoes, and
instruments. During his journey south from latitude 85° N. Lat., F.
Nansen (1897) was also subjected to “attack” by a great number of
arctic foxes which strewed about his belongings. Now, arctic foxes
are significantly more cautious. Nevertheless, factory workers in
Yamal, Taimyr and others have more than once reported the pillage
of the meat and fish stores and other products by arctic foxes. When
an arctic fox is driven with reindeer, the exhausted animal hides
under a fur thrown on the snow. Arctic foxes on Ushishir Island are
very unwary (Kuril chain; Klumov, 1960).

Mousing animals and also migrating arctic foxes frequently no-
tice humans only at a distance of several meters. In this case, the
surprised animal sits down, looks at the person, sometimes stretches
and then jumps up and continues on its way. On the Commander
islands, where arctic foxes are protected, they inhabit villages,
bringing their pups near houses or sheds. The parents, usually in turn,
keep close to their litter, and after becoming accustomed to people,
take the food from their hands. They may wander for hours only a
few steps behind people, barking and sometimes sniffing inhabitants
(Cherskii, 1919—1920).

In captivity, arctic foxes quickly become tame. Getting accus-
tomed to humans, some arctic foxes return after being set free. In the
arctic fox, conditioned reflexes are easily developed. On the Com-
mander islands, the animals knew well, by the clock, the time of their
supplemental feeding and quickly respond to a sound signal. To a
whistle, accompanied by the distribution of dried salmon or fur-seal
meat, “arctic foxes appeared as if from underground, ran immedi-
ately, and yelping jumped on the handout” (Barabash-Nikiforov, 1937).

The relationships of the arctic fox to various animals are diverse.
Above were mentioned reindeer and polar bear. Flocks of brant geese,
gulls and skuas actively repulse the attack of the predator. On the

229 Commander islands, the arctic fox drives pigs away from the flotsam,

326

but at the same time, it plays with tethered goats. Arctic foxes are
afraid of cats and dogs, especially where people incite them. In Novaya
Zemlya, arctic foxes follow dogs to carrion, and sometimes, espe-
cially the young, play with them and even with humans (Shrenk,
1855; Nosilov, 1909; Cherskii, 1919—1920).

On a level surface, a dog can easily catch the arctic fox, but in
hilly or tussocky places or in ice hummocks, the arctic fox runs away
from it. The manner of the arctic fox is similar to that of the corsac
fox; it runs in zigzag with sudden sharp turns; it confuses dogs.

In catching lemmings, the arctic fox waits for a long time for the
appearance of a rodent from a burrow, then jumps and crushes it.
Listening to the squeak or rustle of a lemming, the arctic fox can in
a few seconds excavate the snow to a depth of 1/2 m or more. While
hunting for lemmings, the arctic fox can easily come within gunshot
of a man. A case is known of an arctic fox which, while digging up
small rodents during a snow storm, was caught by a hunter by the tail.
On seeing a gull on a rock on the shore from a distance of 120
meters, an arctic fox not yet molted went on with slow steps, confin-
ing itself to patches of snow; at 15—18 m, it lay and crept on its belly,
and then with a strong jump caught the gull (Chastin, 1939). At the
molting time of geese, arctic foxes hunt them communally, driving
the helpless bird to their comrades; the arctic fox throws the captured
goose on its back and pulls it. It drinks the eggs of large birds through
holes made by the canines.

On the Commander islands, wandering constantly among the
hauling places of fur seals, arctic foxes search for not only dead
animals, but also pull away the young fur seals from lone females
holding them by a unique method, by the lips; they take mainly the
sick and weak (Barabash-Nikiforov, 1937).

The arctic fox eats part of a large prey on the spot, and carries
the remainder to a burrow or buries it. In years with abundant food,
the arctic fox makes a store, collecting some lemmings, fish or large
birds; it buries their bodies in the ground or in sand, firmly tamping
the surface with its nose; it pushes them under stones, ice blocks and
so on. A case is known on Mednyi Island when arctic foxes took
about hundred eggs from the tin box of some sleeping hunters,
burying the greater portion in a /aida. On the same island, they
observed the return of arctic foxes to their stores to eat them (Cherskii,
1919-1920). All the same, a great portion of the hidden food is not
utilized by arctic foxes.

230

327)

The behavior of the arctic fox changes in various years depend-
ing on the abundance of food, prevalent weather and other reasons.
With abundance of lemmings, white hares and willow ptarmigan, the
arctic fox in the Bol’shezemel’sk tundra wanders little, and restricts
itself the whole winter in proximity to its burrows, frequently hiding
in them. When feeding on a second grade food [source], arctic foxes
behave uncomfortably, but nonetheless do not leave summer habitats.
In case of food deficiency, the animals go to the seashore or migrate
in other directions (Skrobov and Kozhevin, 1957). In years of famine,
arctic foxes move close to the winter stations and villages, lose their
prudence and even defend their prey against humans (Commander
islands, Tomsk district) (Cherskii, 1919—1920; Likhachev, 1930).

The arctic fox takes to water and during migration they swim
across large lakes and rivers, for example, the lower Taimyr, 400 m
in width and a current velocity of 3.5 m/s. They swim across' the
lower Khatanga (1—1.5 km) as well as Yenisei and Ob’, straits in the
Bol’shezemel’sk tundra (2 km wide) and to Kil’din Island (Tret’ yakov,
1871; Formozov, 1929; Rutilevskii, 1939; Shastin, 1939; Chirkova,
1955 and others). In Alaska, the arctic fox has mastered straits up to
4 km wide.

The speed of the arctic fox in water is 22.5 km/h, slower than
in the dog. While swimming, the animal holds its back and tail above
water, the latter at a small angle. When the tail gets wet, indicating
fatigue, the fox quickly drowns (Shastin, 1939). The arctic fox swims
also across rivers containing slush at water temperature of about 0°C;
on being turned around several times in a whirlpool, the arctic fox
does not become disoriented and swims in a defined direction. On
coming out on shore, it shakes itself and immediately runs rapidly
away from the shore.

At the beginning of migration, masses of arctic foxes gather by
rivers and run at the very edge of the water (see also p. 330). As long
as ice is smooth, arctic foxes gather on the lakes, playing and harry-
ing one another. At the sound or sight of an airplane flying at a height
of 100—200 т, the arctic fox takes a defensive pose and lies on its
back (V.P. Makridin); at such a time, the red fox runs away.

On the Commander and Solovetskii islands, according to the
behavior and hunting habits, arctic foxes are divisible into the follow-
ing groups: 1) Sedentary animals, leading an inactive way of life, and
living in permanent places near human settlements, chiefly by scav-
enging; 2) half-sedentary animals, carrying on in a region of permanent

328

habitation, but sometimes moving to places inhabited by neighboring
groups; 3) wandering arctic foxes. On Mednyi Island, the more alert
and active are the tundra arctic foxes who hold aloof from people. On
the Solovetskii Islands, vagrancy is developed mainly among males.
The predominance of males among the migrating animals is, appar-
ently, characteristic of arctic foxes of the mainland tundras
(Barabash-Nikiforov, 1937a; Boitsov, 1937; Chirkova et al., 1959).

The behavior of the arctic fox varies sharply with the beginning
of the reproductive period (Cherskii, 1919—1920; Barabash-Nikiforov,
1937a; Boitsov, 1937; Chastin, 1939; A. Romanov, 1941 and others).
On Mednyi Island, the premating roll-call among males starts early
in January. At night in February, the plaintive howling, yelping, and
squealing of females are heard. In March males frequently urinate
near burrows and feeding places; females urinate, raising the tail, and
allowing males to approach them. The animals run, often holding
something in their teeth—a bone, stick or lump of snow. Playing
between mated pairs alternates with hard fights among males. In the
location of the fight, bits of wool, and sometimes blood are left; on
Mednyi Island, it was observed that at that time, arctic foxes injure
the genital organs of each other. In fights, up to ten animals are
involved. In the northern Taimyr peninsula, nuptial games were ob-
served on 20 April 1936. The mating of arctic foxes is accompanied
by loud screeches.

In some instances, male and female become strongly attached to
each other in the breeding period. On Mednyi Island, a female which
had been taken from her chosen male and kept in an open-air cage
with another male, lived with him the whole spring, but did not mate
with him. During the same period, she always expressed her attention
to the first male, who sometimes passed by. The latter, although he
had his own mate, brought food to the caged female.

During pregnancy the males constantly offer food to their fe-
males and with special diligence not long before the appearance of
the newborn. On Mednyi Island, a male carried fish “myagon’ku”
(Cyclopterichtys venticosus) to a female that happened to fall captive.

Shortly before parturition, the behavior of a captured female
becomes more quiet and cautious. With the appearance of the babies,
both parents feed the young in turn, bringing them food and playing
with them, but most often the mother remains near the burrow to
guard the litter. In case of danger, the female sometimes hides in the
burrows together with the young while the male strives to drive away

231

329

the person, howling and gradually moving away from the litter. In the
absence of adults, the young allow a human to approach within sev-
eral paces, and even after fleeing to the burrow, they appear again
after a few minutes. At the age of 3—4 weeks, the animals leave the
burrow, on Mednyi Island already at the end of June and the middle—
end of July, they start playing independently. Making excursions,
they frequently change burrows. At this time, their barking is first
heard. In Lena-Khatanga territory, the young arctic foxes start to play
within 5 days after leaving the burrow, and within 10 days, they
attempt to catch lemmings and excavate their burrows and begin to
feed on meat. At the age of 1.5 months, they shift over to meat and
other food. In the second half of July, they share with their parents
in hunting molting geese. The mother feeds the young on milk for one
to 2.5 months. 2.5-3 months after birth of the young, the father
abandons the litter. At the end of their life together, the mother also
leaves the litter for a whole day. After the female finally abandons
the young, the litter breaks up. A case is known when, after the death
of the father of one litter, the mother transferred the young to the
burrow of another arctic fox, where both mothers fed the young
together (П’та, 1949—1950).

The more the year is deficient in food, the longer the time taken
by the parents in search for food, and the earlier they leave the young.
With food abundance, members of the family stay together until
autumn, and separate gradually; individual arctic foxes live near
burrows throughout the entire winter. In the former Kolymsk region,
a female was found in a burrow together with her adult offspring,
even in winter (Iokhel’son, 1898).

Seasonal migrations and transgressions. The arctic fox is a typi-
cal migrant, and wandering is of great significance in its life. The
migration of arctic foxes must be understood as a historically estab-
lished phenomenon. If judged by Pleistocene—Holocene remains, this
phenomenon has existed for many thousands of years (Tugarinov,
1927; Gromov, 1928; Kirillov, 1950; Pidoplichko, 1951; Vereshchagin,
1953, 1959, 1960; Serzhanin, 1961).

The reason and stimulus for commencement of migration is in-
sufficient food. This is confirmed by the appearance of massive and
long-distance shifts of arctic foxes in years of the disappearance of
the main food—lemmings. At the same time, the relationship between
migration and food abundance is not invariant. The start of arctic fox
movement away from the denning region is sometimes observed in

293

330

the presence of food. Migration is subject to instinct. In old animals,
migration is also modified by experience. One may assume that the
old arctic foxes react to signals announcing the coming of famine—
an early cold snap with rain and wet snow, early departure of birds,
the appearance of onshore and changeable winds and, possibly, other
unknown factors. It was repeatedly observed that periodic migrations
of arctic foxes from places of permanent habitation are repeated after
2—4 years.

Migrations of arctic foxes start in August-September, depending
upon the time when food disappears in the main breeding places. The
massive movement of animals when the arctic foxes “come in crowds”,
as it is said in the North, occurs in October-November.

The route of arctic foxes before the freezing period runs along
the water’s edge, where a trampled path is formed. At Serebryanyi
[Silver] bay on the western coast of the northern island of Novaya
Zemlya, the whole shore was spotted with tracks of arctic foxes in
1887 and the beaten tracks of animals from all sides met here. On the
sands of the Khatanga river, in 1931, there was a path several meters
wide (Nosilov, 1909; A.F. Chirkova). Part of the arctic foxes move
on a wider front. In the northwestern Taimyr peninsula, arctic foxes
move along in a 2-3 km deep strip along the shore. In the northeast-
ern Bol’shezemel’sk tundra, the animals move along a front of tens
of kilometers, but the majority of them are concentrated in the coastal
zone of Baidaratsk bay (Kirpichnikov, 1939; Chirkova, 1955).

In separate places, either latitudinal—western or eastern—direc-
tions predominate, as in northern Arkhangel’sk district, or meridional
ones, as the well-known route of the coastal arctic fox along the
Yenisei river. Migrations with departure from the tundra to the forest
zone are more noticeable, and early began to attract to themselves the
attention of investigators. Meridional migrations are noticeable only
when they are massive phenomena. In the neighborhood of Taimyr
lake, the autumn migration in 1947 was in a southeastward direction;
the arctic foxes swam across the lower Taimyr from the western
shore (Sdobnikov, 1958).

Animals penetrate the taiga zone in great numbers, going along
the valleys of large rivers or coming out onto open watersheds; from
the sea coast, they move out onto the ice and to the islands of the
Arctic Ocean. The arrival of arctic foxes into the forest zone is
frequently preceded by the flight of snowy owls [Nyctea scandiaca]

331

which more quickly respond to deficiency of food in the tundra and
vacate it earlier.

It has been observed that arctic foxes move as though in waves.
Such migrations of arctic fox groups were observed in Lena-Khatanga
territory. In the interior of the Bol’shezemel’sk tundra, in 1933/34, a
movement from east to west began from beyond the Polar Urals,
which was noticed at the end of October when in the course of a
month there was a migration of well-fed arctic foxes. After a pause
at the end of December, a smaller group of animals moved, and from
January 20th, during the rest of the month, individual animals were
seen running by on three days. The second and third groups of ani-
mals were emaciated. On the sea coast, arctic foxes at the end of
October moved the whole winter in the same western direction, the
first strong wave occurring at the end of November, the next—at the
end of December, and finally—in the middle of February and from
the end of this month until the middle of March and later. The rut,
which took place at the end of March and in April, did not stop the
movement of arctic foxes, which continued running in the same di-
rection as in winter (A. Romanov, 1941; N. Dement’ev, 1955).

In winter, arctic foxes spread out over a huge territory of tundra,
forest and ice landscapes. The total area inhabited by the species
increases at this time by several times and at the same time popula-
tion density in the tundra decreases. The majority of arctic foxes die
in unaccustomed life situations, from hunger or diseases, and are
caught by hunters. Long-distance migration of arctic foxes in the
majority of cases is “a way to death”.

Those animals remaining alive in February—March, sometimes
even in January, migrate back; in some years this extends until April
and May. The first tracks of arctic foxes from the south appeared
near Taimyr lake on the 20th of March, and the movement from the
seashore into the depths of the tundra in 1948—1949 ended by the
middle of May (Sdobnikov, 1958). Apparently, only a negligible
portion of the arctic foxes return to the tundra. Individual arctic foxes
remain during the warm period in uncharacteristic types of land-
scapes like forests and on the ocean ice.

Long-distance migrations are more often carried out by males (on
the Solovetsk islands, up to 80%), but cases of long movements are
also known in females, for instance in 1928, to the neighborhood of
Krasnoyarsk. Long-distance movements seriously deplete arctic foxes

8и1рээ14 ay} JO зэмерипод и1э410$—6 ‘1еэА oY} JO SOUIT} зпомел UI 9d1 UO злэзипоэиэ рие JOJUIM ит риеритеш oY) UO ззиэшэлош эоие}$1р-8ио]
JeNplAIpul—zZ ‘зиоцелаи 1эдим рие ицицие JO иопоэир—1 :(влодичо `4`У) SOXOJ эполе JO ззиэшэлош ээиезтр-Зио] рие зэтол UOneISI “Lg ‘914 ZEZ

008 06

1 05

р 051 ou
SESS
7

о&х

353

and suppresses their reproductive capacity (Sluchevskii, 1886; Skalon,
1928; V.K., 1929; Boitsov, 1937; Chirkova, 1955, 1959 and others).

In the forest zone, arctic foxes were observed and frequently
hunted at 1000—1800 km, and in exceptional cases, 2000 km (for
example near Komsomol’-na-Amur), and in the Arctic Ocean, at
1000 km from the limits of summer habitation. The average daily
route encompassed by the animal is, apparently, about 20—30 km.

At the end of the 1930’s, 172 arctic foxes were marked in the
Novyi Port region on Yamal, of which 31 animals were re-captured
(Sdobnikov, 1940; Fig. 88). A maximum translocation of about 1200
km in a straight line as it went across ocean ice to the north island
of Novaya Zemlya, was made by one animal. The majority of arctic
foxes moved off in a southwesterly direction; the farthest ones were
caught near the village of Muzhi on the Ob’, at the mouth of the
Pechora, at the Snopa river on Chesh Bay, at the southern end of
Novaya Zemlya, and at its northern limit.

When food decreases in years of the animal’s abundance, almost
all arctic foxes migrate or disperse to different areas away from their
denning places. Districts to which they dispersed to the south for the
majority of migrating arctic foxes were to be found in the northern
subzone of the forest. When a peak of migration is reached along a
permanent route, one may observe large numbers of foxes. Thus, in
1887 during one stroll on the western shore of Novaya Zemlya,
Nosilov (1909) encountered hundreds of animals, and in the course
of two weeks in October, by his estimate thousands of foxes passed
by. After migration to the southern island, only single individuals
remained. On the southern coast of Belyi Island at the end of Septem-
ber 1930, tens of foxes were seen, through binoculars, wandering
about awaiting the formation of fast ice (A.F. Chirkova). On Dikson
Island, on good “movement” days 5—6 arctic foxes were seen directly
in a distance of 1 km (Heptner, 1936). On the lower Taimyr river,
between 18 and 20 September 1935, each day up to 21 foxes swam
across while on September 1947, 10 to 15 foxes were observed at one
time within a radius of 2 km in this region (Sdobnikov, 1958). On the
Chelyusk peninsula, during the time of the arctic foxes passing at the
end of October, one might see at one time up to 46 animals, after
which the number dropped sharply; during winter, there were only
separate individuals. The total number of foxes that passed through
an area of 10 km in autumn 1932 constituted not less than 1500 to
2000 individuals, and in spring, during the period of their return,

233

334

—— =

a
[7

I
Е

NY
\]

Fig. 88. Directions and limits of migrations of individual arctic foxes marked on
Yamal (from Sdobnikov, 1940): 1—direction of movements; 2—no movements.

there were not less than 800 (Rutilevskii, 1939). On the right bank of
the lower Khatanga, upstream from Cape Bol’shaya Korga, tens of
passing arctic foxes were counted each day on 24 and 25 August
1931. Along the left bank of this river near the mouth of the Novaya
river, from 2 to 80 fresh tracks of arctic foxes were noted on a 10-
km route at the beginning of October, and below Khatangsk village,
it was possible to observe 2 to 4 migrating arctic foxes at one time
at any time of day starting from September 17; after ice formation on
October 10, arctic foxes disappeared. In that period, no less than 1000
animals passed by Khatangsk village (A.F. Chirkova).

In Arkhangel’sk district, judging by the number of prepared skins,
the number of arctic foxes emigrating for long distances only in a
south and southwestern direction constituted, in some years, about 10,
20, or 30%. No less than this number of arctic foxes departed to
the north across the ocean ice. In this manner, during years of exten-
sive migration up to 2/3 of all arctic foxes abandon the tundra zone.

At the time of massive movements, the behavior of arctic foxes
changes. At the beginning of migration, the direction of movement of

235

335

individual animals is variable, undefined and might be directly орро-
site one another. In this time, arctic foxes wander nervously and swim
across rivers and canals. Coming to the banks of large and as yet
unfrozen rivers, groups of arctic foxes start to change direction along
them, running along the sea coasts and along mountain ranges and
valleys; they move in response to the density and depth of snow
cover, the weather, especially winds, and also to the availability of
food along the way.

At the beginning of migration, arctic foxes move in pre-twilight
or early morning accustoming themselves to the trails trodden by
previous confreres; without looking aside they hurriedly overcome
the difficulties along the way. Later arctic foxes energetically run,
trot or even gallop day and night, with this frequently barking or
howling. Being distracted for a short time to search for food, their
hunger barely satisfied, the animals again start on their predetermined
way.

On clear moonlit nights or during the period of the northern lights
[aurora borealis], active movement is initiated. On their route, the
arctic foxes move through villages, especially at night, and it is only
during the light part of the 24-hour period, when people and dogs are
present on the shore that the animals detour or lie in wait for the
departure of people. Their instinct for wandering is so strong that the
animals may stop for some time only to spread out at the beginning
of migration to feed up, although the animals still remain in the
denning region or have only started to migrate. During the peak of
their movement, hunting for food detains the animals only briefly.

Arctic foxes from different geographic populations may sometimes
mix during migration. Thus, in the winter of 1956/57, in a year of migra-
tion from the east, an average of 50% of the arctic foxes captured in the
vicinity of Kara at the very edge of the Bol’shezemel’sk and Yamal
tundras were related by their fur to the Obdorsk type, characteristic of the
Yamal foxes. In the western Bol’shezemel’sk tundra, such animals соп-
stituted 30%, and on the Kanin peninsula, less than 10%; all those
remaining were referable to the local Pechora type characteristic of the
European north (Chirkova et al., 1959; Fig. 82).

Reddening of the fur that is observed in arctic foxes and
other carnivores on the underside of the paws, rarely in the axillary

In microscopic view, the reddish color stains equally every hair. The color is
not soluble either in water or in soap, and it resists the action of alcohol, ether,
chloroform, xylol and hydrogen peroxide (Chirkova et al., 1959; Shilyaeva, 1961; А.
Solntsev).

336

depressions and on the belly, is obviously caused by skin gland se-
cretions connected with periods of great stress on the organism, and
appears particularly intensely during the time of the animal’s migra-
tion. Thus during the extensive migration of arctic foxes in the
northeastern Bol’shezemel’sk tundra in the autumn and winter of
1951/1956, the reddening of the fur of the foot pads and between the
toes was noted in approximately half of the arctic foxes captured in
the Kara region (42% bright color). In the following year during the
absence of migration, reddening of the paw fur was only noted in a
small number of animals (2% bright color in all) (Chirkova et al.,
1959).

Basic migration routes and long-distance movements of arctic
foxes are known over a large part of the species’ range (Fig. 87). On
the Kola peninsula, the arctic fox migrates mainly along the valleys
of Kamenka, Kochkovka, and other rivers, [flowing] from the interior
of the peninsula to the sea. From the northwestern Kola peninsula it
runs southward to Murmansk and farther, along the railway line.

Rare cases of the appearance of individual foxes were observed
in the territory of the Lapland preserve and on the Tersk shore in the
Olenitsa and Kuzomen’ regions (Ozeretskovskii, 1904; Dubrovskii,
1939; Zolotov, 1904; Nasimovich et al., 1948; Drebentsov, 1959).

After the formation of fast shore ice and the appearance of ice
at the mouth [entrance]* of the White Sea, the arctic fox moves off
to the Kanin side, from whence they return in March, before the start
of ice breakup. The large exodus of the arctic fox is repeated every
4 to 6 years. The emigration of arctic foxes to the Varanger Fiord
side, and the event of their appearance in northwestern Finland near
Enontekléssaé is notable (Kleemola [and] Laine, 1961).

Counter-movements across a wide distance take place in the
European north, to the east of the White Sea, from the northeast to
the southwest and west through the Bol’shezemel’sk, Malozemel’sk
and Timansk tundras (Folitarek and Chirkova, 1930; Sergeev, 1934;
A. Korsakov, 1941; G. Naumov, 1941; Maksimov, 1945; Chirkova,
1955). Arctic foxes skirt to the south of Chesh, cut across the Kanin
peninsula or follow around it, turning to the north along the eastern
shore, and drop down to the south along the western; they migrate
along the Zimnii coast of the White Sea to Dvina Bay. The arctic fox

*The Russian word used is gorla, meaning throat—Sci. Ed.

236

337

frequently crosses the lower Pechora and other large rivers in specific
places.

Many arctic foxes migrate southwards along the Pechora, Mezena,
Pinega, Northern Dvina, Onega rivers and their tributaries to the
upper Vychegda, Sukhona, Yuga and others. The most extensive
emigrations of the arctic fox, up to 58—56° М. Lat., were known even
in the past century, and also in the present century in southeastern
Latvian SSR, in Pskov, Novgorod, Leningrad, Kalinin, Vologda and
Kostroma districts, and in the northwest and east of the Kirov district.
Migrations were noted as far as the vicinity of Troitsko-Pechorsk
village and the mouth of the Ilych river, and also in Ust’ Kulomsk,
former Storozhevsk and Letskii regions (60° N. Lat.) of the Komi
ASSR (Middendorf, 1869; Sluchevskii, 1886; Dmitrieva-Sulima, 1911;
Paramonov, 1929; Ognev, 1931; Lavrov, 1932, Sergeev, 1934; Guber,
1937; Ostroumov, 1949; S.V. Shibanov, A.F. Chirkova).

In the Pechoro-Ilych preserve, arctic foxes during the period
from 1929 to 1949 appeared in the course of 9 winters: in 1929/30,
1935/36 and 1937/38 there were massive migrations, and in 1933/34,
1938/39, and in the course of four winters from 1944/45 to 1947/48,
individual animals were encountered. Apparently, movement of the
arctic fox on the territory of the preserve was limited by the depth of
snow cover; in years of large transgressions, the average maximum
depth of snow cover was 74 cm, but, in winters of occasional appear-
ances—98 cm. In heavy snow years, the arctic fox often remained
along rivers, moving on the ice, being malnourished and diseased
(1929/30 and 1935/36). In light snow winters (1937/38) they moved
freely through the taiga. There is a supposition that occasional arctic
foxes, moving into the preserve in 1944—1945 continued to live there
and did not depart for 4 years; after the capture of two animals in
1947/48, no more foxes were encountered (Teplov, 1960).

Some of the arctic foxes from the northeastern Bol’shezemel’sk
tundra move over to Vaigach and Novaya Zemlya, across the ice of
the Barents Sea, and then arrive at Kolguev [island]. The migration
of arctic foxes in Novaya Zemlya was intensive in the last century
along the western and eastern shores of both islands (Nosilov; 1909).
Good passages of arctic foxes on Novaya Zemlya occur twice a
year—in autumn and spring (Dubrovskii, 1941). In different years,
arctic foxes move from Bol’shezemel’sk tundra to the Urals., Waves
of massive migrations in the north of Arkhangel’sk district were
noted in the winters of 1909/10, 1914, 1926/27, 1929/30, 1935/36,

338

1937/38, 1944/45, 1946/47, 1947/48, 1954/55, 1956/57 (Dmitrieva-
Sulima, 1911; Folitarek and Chirkova, 1930; Bashkin, 1940;
A. Korsakov, 1941; G. Naumov, 1941; Maksimov, 1945; Chirkova,
1955; Chirkova et al., 1959).

Basic migration routes on the Yamal peninsula, when, accord-
ing to Nentsi stories, foxes run “in herds,” go in a meridional
direction, along the shores of Ob’ Bay and the Kara Sea, and also
along the central Khoi watershed. The arctic foxes move out over
a wide front along the ice of the Kara Sea.

Less significant are migrations along the Taz and Pur rivers to
the east on the Yenisei. The most southern places in west Siberia
where arctic foxes appear are noted in the vicinity of Nyaksimvol’
(63° N. Lat.); in the belt of southern taiga along the course of the
Irtysh and Tobol, below Tobolsk and even Yalutorovsk (56.5° N.
Lat.), below Tevriz (57.5° N. Lat.). Dispersal of foxes to the vi-
cinity of Tom’sk (57° N. Lat.) is known (Skalon, 1928; Formozov,
1935; Andreev, 1937; Laptev, 1958; M.A. Sergeev). Long-distance
movements of arctic foxes into the forest zone of west Siberia
took place in 1921, and in the seasons of 1926/27, 1929/30, 1932/
33, 1937/38, 1941/42, 1944/45, 1947/48, 1954/55, and less
significantly in 1930/31, 1933/34, 1935/36, 1939/40, 1946/47
and later. Migrations of the arctic fox are also known in 1910 and
1913.

In northern Krasnoyarsk territory, the arctic fox takes itself
upstream along the Yenisei to the mouths of Lower and even the
Podkamennaya [stony] Tungusk. Occasional arctic foxes reach
Krasnoyarsk (V.K., 1929, and others), the upper course of Lower
and Podkamennaya Tungusk (60° N. Lat.), to the region of the city
of Bratsk, even the villages of Aban and Taishet along the main
Trans-Siberian railway (56—57° М. Lat.). From the Taimyr peninsula,
large numbers of arctic foxes migrate along the Pyasina and Nizhnaya
Taimyr rivers and also along the shore of Taimyr lake, departing from
the tundra zone through the belt of northern taiga forest to the Central
Siberian Plateau. A well-defined migration of arctic foxes takes
place along both shores of the Khatanga gulf and Khatanga river
and its tributaries, the Popigai, Kheta and Kotui, where the arctic
fox reaches the upper courses of these rivers. To the north, arctic
foxes disperse across the ice to all the nearby islands, in particular
Severnaya Zemlya and Vize island—at 78° N. Lat.

237

339

Large migrations and long-distance dispersal of arctic foxes in
the northern Krasnoyarsk territory are known in 1860, 1913, 1922/
23, 1925/26, 1927/28, 1931/32, 1935/36, 1944/45, 1946/47, 1947/48,
and 1954/55 (Tret’akov, 1871; Aleksandrov, 1927; Buryakov, 1929;
N. Naumov, 1930; Heptner, 1936; Shastin, 1936, 1939; L. Popov,
1939; Chirkova, 1955).

Arctic foxes in Yakutiya wander not only along the rivers but also
in the interfluve country along the Kharaulakh, Chekanovsk and other
ranges. Transgressions were observed in the upper Anabar, Olenek,
Vilyui, and Lena rivers (Mukhtuya and Olekminsk villages, 61° N. Lat.)
and along the Vitim (Mama village, Irkutsk district, 58° N. Lat.), in the
vicinity of Yakutsk and Krest-Khal’dzhai village on the Aldan. Arctic
foxes were found along the Omoloi and Yana rivers to the latitude of
Verkhoyansk, along Indigirka to Oimyakon village (63° N. Lat.), on the
upper Kolyma to Seimchan (62.5° N. Lat.) and its tributaries, the
Yasachnaya and Korkodon (Tkachenko, 1932; Tugarinov et al., 1934;
Kolyushev, 1936; Guber, 1939; Skalon, 1940; Skalon et al., 1941; A.
Romanov, 1941; Belyk, 1953). Extensive migrations of foxes take place
here in the tundra zone and along the shores of the Laptev and East
Siberian seas, in both easterly and westerly directions. Arctic foxes
come to the Anabar river basin following the migration of wild rein-
deer herds from the eastern Taimyr.

Commonly, arctic foxes disperse across the ice of the Arctic
Sea, to the New Siberian and other islands (V.U., 1935, Zarovnyaev,
1933; A. Romanov, 1941, and others). Masses of arctic foxes,
together with seals and polar bears, remain in winter among the
[ice] hummocks in the sea 250 to 350 km from the mouth of the
Olenek river; here these animals seek their living. After strong
winds and snowstorms, the arctic fox wanders back to the sea-
coast (Vasipev, 1935).

Migrations of arctic foxes in the western and eastern regions
of the Yakutsk ASSR do not proceed simultaneously. Large migra-
tions of arctic foxes to the east of the Lena (along the Kolyma,
Indigirka, Alazeya and others) were observed in 1895, 1903 and
1944/45; and in the western regions, in 1926/27, 1932/33, 1947/48,
and 1954/55. In 1929/30 and 1935/36, long migration of arctic foxes
originated, apparently, on the whole territory of Yakutiya (Iokhel’son,
1898; Buturlin, 1913, E. Sh., 1930; Tugarinov et al., 1934; Skalon,
1940; A. Romanov, 1941; Chirkova, 1955).

238

340

Е. Vrangel’ (1841) even saw arctic foxes on the ice of the East
Siberian Sea between the Medvezhie islands and the De Long
strait. Some information is available on movements of arctic foxes
along the shores of Chaun bay to Wrangel’ island and back at Cape
Uelen, and eastward across the Bering Strait, and also in a south-
western direction along the shore of the Bering Sea (Mineev, 1935;
Chirkova, 1955 and other materials).

_ Transgressions of arctic foxes are rarely observed оп
Kamchatka—in the Parapol valley region on the isthmus, on the
western coast to Ust’-Bol’sheretsk, the southern end of the penin-
sula and the northern Kuri! Islands; on the eastern coast of the
peninsula—to Ust’-Kamchatsk and the Kronotsk peninsula. In 1895
and 1896, several arctic foxes were caught near Petropavlovsk
(Paramonov, 1929; Sergeev, 1936; Guber, 1939; Yu.V. Averin).

Arctic foxes were noted at Gizhiga and Penzhino (Ognev, 1926)
and along the western shore of the Okhotsk Sea (Sergeev, 1936;
Guber, 1939). Nordmann (1861) was the first to point out the oc-
currence of the arctic fox in the lower Amur. Later transgressions,
or possibly the transporting of separate individuals on ice, were
observed at 51—52° М. Lat. on the western shore of the Tatar strait,
in the Ul’chsk region in the winter of 1949/50 and in the vicinity of
Komsomol’sk-on-Amur in the autumn of 1958 (Belyaev, 1959).

F. Nansen encountered arctic foxes on the ocean ice below 85°
20’ N. Lat. in April 1895, 300 miles from the nearest land (Franz Josef
Land). Brusilov (1934) during the period of the “Northern Anny” drift
[ship in ice] noted the presence of arctic foxes in the winter of 1913/
14 on the ice between the islands of Novaya Zemlya, Franz Josef Land
and Severnaya Zemlya, approximately from 77° to 82° N. Lat. and from
60° to 80° E. Long. De Long (1936) saw arctic foxes and their tracks
at a latitude of 72° to 77° N. Lat. between Wrangel’, Geral’d and Bennet
islands. During the voyage of the ice-breaker “С. Sedov” and later on
the drifting polar stations “Severnyi Polyus” [North Pole] 2, 3 and 4
in the “inaccessible pole” and other parts of the ocean, arctic foxes
were found between 77°-87° 45’ М. Lat. and 129° Е. Long—157° W.
Long. at distances ranging from some hundreds to 1000 km to the north
and northeast of the New Siberian islands (Badigin, 1940; Buinitskii,
1945, 1946; Karelin, 1945; Ostrekhin, 1945; Chapskii, 1946; Rutilevski
and Uspenskii, 1957; Kirillov, 1958). The most northern point at which
the arctic fox has been observed is ~88° N. Lat., at a distance of 800
km from the nearest land—Ellesmere Land (Uspenskii, 1956). The

341

majority of occurrences of arctic foxes met with on ice are noted in
November—December, and rarely in March—April.

In the Western Hemisphere, arctic foxes migrate also in various
directions, both to the south and to the ice of the Arctic Ocean. On the
‚ eastern Atlantic coast of the North American continent, arctic foxes
descend along the treeless regions significantly farther south than on
the western Pacific [coast]. Up to 90% of the arctic foxes within the
arctic archipelago spend the winter on ice. Arctic foxes even disperse
from Canada to Greenland.

Reproduction. Animals begin to reproduce at the end of the
first year of their life, although they attain full maturity in the
second year. Development of the reproductive organs begins with
increase of the testes in autumn. In a year with unfavorable con-
ditions, the formation of spermatozoa and their appearance in the
excretory ducts in adult wild arctic foxes of Yakutiya, began in
February at a testes weight of 2600 mg and more. In young 9—10-
month-old males, sperm was formed a month later (N. Sokolov,
1957). Decrease in the volume and loss of elasticity of the testes
in arctic foxes on farms takes place at the end of April-beginning
of May when the prostate gland ceases its activity. The male
produces sperm during the course of 2—3 months. The testes of
arctic foxes are larger in size compared to those of red foxes and
the ejaculate contains a large number of spermatozoa. Among
captives, the arctic foxes with the least weight in summer but
which rapidly increased in autumn, were found to be the best re-
producers.

In the course of the winter, ovaries and oviducts of females in
the pre-estrus condition develop earlier in adults than in young.

Ovulation of the spontaneous type does not depend on the time
of mating. Before the beginning of ovulation, the ovary of the arctic
fox is larger than that of the red fox, and although the ova are
smaller, the number of Graafian follicles is larger; this fact deter-
mines the fertility of this species.

Estrus in arctic foxes is more prolonged than in the red fox, and
on animal sovkhoz [state farms], it lasts 12—14 days, but under
natural conditions the female remains [in estrus] for only 4—5 days

S8Middendorf, 1869; Cherskii, 1919—1920; Paramonov, 1929; Barabash-
Nikiforov, 1937a; Dubrovskii, 1940; A. Romanov, 1941; Rakhmanin, 1959;
Skrobov, 1958; Chirkova et al., 1959; Starkov, 1932, 1937, 1940; Tavrovskii,
1946; Il’ina, 1952 and others.

2539

342

on the average. During estrus, the mammary glands enlarge to 0.5
cm in diameter. In the copulatory period, the male and female pair;
on farms this may last from one minute to several (4) hours (Boitsov,
1937); coitus may occur repeatedly.

In nature, the breeding season extends from the second half of
winter to the end of summer. With the appearance of the first rays
of the sun, courtship behavior begins. At this time, 1-2 males, rarely
more, run after one female (in captivity, 2—3). The peak of rut and
mating are often observed in March—beginning of April. Adult ani-
mals commence breeding first, and then the young (on Yamal,
10—15 days later), and last are those which have returned from
migration.

In various regions of the USSR, over a wide area the breeding
dates of arctic foxes are nearly the same (Table 28), but the ap-
pearance of young in the northern subzones of the tundra is normally
2—3 weeks later compared to the southern (Yamal, Taimyr).

In one or another place, the breeding season and fertility of the
arctic fox can change annually in connection with the dynamics of
nutritional conditions and age composition of the populations. The
mating period in one place fluctuates within 2—3.5 months, and the
time of birth of young arctic foxes is shifted by 1.5—2.5 months on
the continent and up to 3 months in the Commander islands. Under
favorable conditions, a local concerted mass rut is observed; in a
year following inadequate food, it is undetectable, prolonged, or
greatly delayed. In certain years, fertilization and implantation occur
in 70-80% of the females and, in some places, possibly even more.
At this time all littering burrows are occupied by young, and some
arctic foxes deliver their newborn under clumps of grass. In other
years, the majority of females remain barren and 10—15% or less of
the littering burrows are occupied (see also р. 365ff.).

After mating, arctic foxes explore the region which they have
chosen, and the female begins clearing the burrows. This takes
place at various times of day depending on the general course of
reproduction, mainly in connection with the nutritional conditions of
the year. In the Solovetsk islands, arctic foxes began to choose the
place for the future nest but the burrows were ready 5—15 days
before parturition. One female which was bred on 19 April, occu-
pied a burrow together with a male on 21 May, and gave birth on
6 June. In Yamal, the burrows are also cleared from March to the
beginning of April—and on Belyi Island, to 20 April. In the lower

343

Yenisei, already in March, a female wandered around a hillock
which she chose for her future lair, making a round of up to 5 km.
In the northwestern Taimyr peninsula the clearing of the burrow
progressed from the second half of April, and in Lena-Khatanga
territory—the middle of March; a pregnant female littered here on
7 June, and emerged after parturition on 10 June. On Mednyi
island, the Arctic foxes now distribute themselves among the bur-
rows after mating.

In captivity, gestation continues 49—56 days, 52 on the average
(also confirmed by observations in nature). In the earliest and latest
parturition (45th, 62nd* day), newborn often die. The abdomen
obviously increases in size 1—2 weeks before pupping, sometimes
earlier. The female usually whelps in a burrow, but sometimes
directly at the entrance: this is often a young female that did not
prepare the nest, and later, are carrying the newborn to the burrow.
The duration of parturition depends upon litter size, after approxi-.
mately half an hour, the first pup appears; sometimes, parturition is
extended over 2—3 days. Most often, the female delivers herself of
her litter** at night or early in the morning.

In the tundra, mass appearance of the newborn arctic foxes
occurs in May and June and coincides with fawning of reindeer, the
return of willow ptarmigan, and the arrival of geese, gulls and other
birds. The earliest parturitions of arctic foxes known are mid-April
on the Commander islands, the end of April on Novaya Zemlya and
Belyi Island, and the very latest, in June in Severnyi (North) Yamal,
Lena-Khatanga territory, Commander islands, and other places.

March April

Number of matings

Five-day interval
HE

Fig. 89. Times of mating of arctic foxes in island husbandry on Kil’din,
Murman (from Zolotov, 1940).

*Appears inconsistent with previous values for length of gestation—Sci. Ed.
**In Russian original, literally “burden’—Sci. Ed.

eS
8561 UAsiIds ‘мм
‘AOYIUQOPS ‘6661 ‘ulURW

$961 "5561 “BAO = -YABY 6661 8S6l
0561 ‘влодичо = WYO 1961 ‘влодичо ‘л040195
—6r61 “PUL IT ‘1y6 “Aouewoy ‘у ‘OP6l ‘ИУ$ "5561 “BAO 9761
1561 ‘^010} ‘лоиешоз “у "6561 -^о19па ‘6561 -ХИЧО :6561 “TEYSAOIAR L
“DIIN-Yseqeieg $961 6561 “IIS ‘и13$8Ч$ 6561 “JoqnD 8861 ‘лэдиэшэ 1561 ‘ИУ$лол "6561 “HIS
‘0761-6161 “BAOWMYD -AOIARL 6981 ‘1э4по -9¢6] “UINIAL ‘9€61 “1561 “aNd ‘6061 -^о49па :6861
‘1151940 «= “SEH ‘лээи ‘рмориэррид ‘AgysnAjOy ‘Aoysndjoy “AOURQIYS “AO|ISON “AOZOUWMO sIOyNy
ounr jo oun
Suluuisaq 0} jo 8шии3э4
Ави jo —Ави JO
ounr—Aep — jyey puosas — — УЧ puoses — — SUIdjoyM 5$
19181 Аш иг цэлэ Aine
Ajng jo pue Ane pue ounce sung 40 Зиши! 334
SUIUUIZAg 01 jo Зиииэ4 suluulsoq = jo эррии о} ‘Ajqeqoid ‘pue ounr
Ajng 0} [dy sung Ави Jo ИеЧ —oung 0} АИ 0} judy Ави jo Sulu эипГ о} [dy jo Зиши 899 (01—10)
jo эррии шюоз4 ‘Азию pug 151} WOLJ jo pus шолН JO pus шол4 -ш8эа шол4 Jo риэ шол4 —Ав/и Jo pug Suidjay
Judy
jo Зииит8э4а Kew;
pue Чэлви jo 8шиш8эа = ud w—youe yy YyoiejAy Jo
yoey| — jo ИТ puosss yore, АвИЕ-Чоеи ‘па9у ‘yore 40 ЭРРИД Jey puosss и] ЗиЦеш $58/
Aew Judy эип[ jo ounr jo
jo Zuluuisaq Ави jo АИ jo pus 40 эррии Зиии!834 0} Ави 0} suluutsaq 0} Kew;
01 Алепие[ pus 0} Але 0} Чо Кле 0} Ашплаэ4 Алп9э4 jo = Aueniqaqy ю —4^лп19эя jo 40 риэ 0} (оо)
дю риэ woly “niga 014 -п19э4 шоля дю pud шол4 эррии шо24 эррии Wooly s[pplw лол АлеплаэН wolj $1159 Ча

рие]$] .[э8 цел 1214391 р spur|si
Spuejst WM 45510 BAIINAL A e[nsutuad 2119915 —- YS, Josue eA] WZ UIP их pue
Jopuewwiosd UIO}SBIYWION UIOYION 1Ацие] и1э}5эм чом (194340 М еАелом в пзишо4 во

59х0} поле JO $эии) Зирээля '82 эае1,

Ove

—

345

The appearance of litters near burrows was observed from the
end of May until August, and more often from the second half of
June to the beginning of July; it is very rare when young appear
later than August. The appearance of delayed litters in August
(“of a mitten” size) or when still in summer coat (krestovatik)*
in November, is often interpreted as a second litter: Actually, the
late whelping is explainable as delayed estrus or a second estrus
after failure to conceive, abortion, or death of the newborn. Cases
of repeated estrus in captives are known, up to 4—5 times 7—30 days
after the first mating. In some years, 5—8% of the females come
into estrus for a second time. Apparently, the males, in some cases,
retain the capacity to mate with the female in a later period. Thus,
the testes of an arctic fox from Yakutiya, taken from its family at
the end of July, still contained mature sperms (N. Sokolov, 1957).

The emergence of the litters from the burrows takes place in
the course of August—October, depending on nutritional conditions,
and in the unfavorable years (Kara region), even in July. In years
of abundant food, the litters remain near the burrows during the
course of the entire autumn and winter. |

Fertility of the arctic fox is very high for a carnivore. Fertility
can be judged by the number of corpora lutea in the ovaries, of
embryos, and of the remains of the post-partum pigmented spots
[placental scars] in the uterus. The latter are retained until the
beginning of proliferation of the reproductive organs prior to the
next estrus, and sometimes (in starvation) to the new estrus. In
1934, when food was abundant on the Kolyma, intensive reproduc-
tion was proceeding among the arctic foxes. In 27 females studied,
the number of placental scars in the winter 1943/44 was from 3 to
24 (in the majority—10, 11, 15 and 17), averaging 12.7 for each
parturient female (Cha, 1953).

The situation becomes more complicated when the population
decreases. Thus, in the Kara tundra, the summer of 1956 was not
favorable for reproduction of arctic foxes due to the disappearance
of lemmings. Out of 22 adult females investigated in the winter of
1956/57, 8 remained barren (placental scars in uterus absent); 6
females revealed from | to 6 ill-defined scars, and the remainder
from 6 to 16 well-defined scars at the same time (in one even in
the cervical region of the uterus) and up to 10 ill-defined patches

*Local word; see р. 351—Sci. Ed.

346

in each. The latter might have been traces of the placentae of
embryos resorbed before parturition. The largest number of scars
was 21. On the average in each pregnant female about 1/3 of the
embryos did not live until parturition; on the average, of 10.2 em-—
bryos, only 7.1 were likely to have been born (Chirkova et al., 1959).

The greatest number of placental scars found in one arctic fox
on Belyi Island was 26 (Tyulin, 1938). The possibility of one female
nursing such a large number of pups has never been confirmed.
Animal breeders report up to 16 newborn delivered by one mother.
Cases of finding more than 20 young arctic foxes in one burrow are
most frequently explained by the union of two or more litters. Thus,
in the Kolymsk area, when a colony of burrows was excavated, up
to 40 or more pups were found. On the Commander islands, a case
was noted of two females with their pups of various ages living
with a pair of parents with its own litter in one burrow. Here also,
females were observed who, beside their own offspring, had 3—4
foster pups. When breeding arctic foxes in captivity, a maximum
number of 22 embryos was observed, but usually a part of them
was resorbed or the pups were born dead.

Geographic variation in fertility is observed between continental
and maritime arctic fox populations. In continental tundras and on
Novaya Zemlya, great fluctuations are observed in litter size and
the proportion of infertile (unmated) females in the various years.
The number of pups in a litter in the continental tundras varies from
8—12 to 20 and more™ on the average in years with abundant food,
and decreases to 3—5 in years of famine. In the maritime tundras,
for example on the Kola peninsula, in northeastern Siberia, and
especially on the Commander islands, the size of litters (to 10—13
pups) and the number of females participating in reproduction
varies to a lesser extent from year to year. On Kil’din island, poorly
nourished females produced 4—6 pups, and females nourished better
than average—1l1—13. On the Commander islands, reduction was

242 repeatedly observed in the number of young in litters from 6—7 to
2—3 on the average in separate places—“ukhozh”* depending upon
the abundance of natural food and supplemental food. The absence
of large litters is also characteristic for the island husbandry in
North America and was noted in the maritime populations of arctic

“То 22 in Yamal (Rakhmanin, 1959).
*Literally, “pasturage”; in this context, foraging places—Sci. Ed.

347

foxes in Greenland (Lek, 1957)*. Apparently the character of this
variability is conditioned by the greater or lesser constancy of the
supply of food.

Therefore, the fertility of arctic foxes is lowered according to
the number of infertile and unmated females as well as to the death
of offspring—the resorption of litters during the time of embryonic
development and the death of pups after birth. In fox farms, unmated
female averaged 7% at the end of the 1930’s and those which
were infertile—15%. Embryonic mortality attained 36%.

The sex ratio in populations of the arctic fox is almost | : 1, but
this ratio may change depending on season, food conditions of the
year and the age composition of the population. In the majority of
cases observed, males predominate. In Kola and Saltykov fur
sovkhozes [state farms], 2—3% fewer females were born. In the
latter, during the period 1950—1957, deviations established a pre-
dominance of males in one year in one sector of the sovkhoz up to
4.4% (of 1100 pups) and the dependence of sex [in litters] on the
age of females; in the litters of young (up to 2 years) and the old
(6—8 years) females, males predominated (51—54%), while in those
of moderate-age—females (51—51.5%). An increased number of
males was observed in large litters (Ivanter, 1962).

On Yamal, at the beginning of the 1930’s, 50.5% males were
caught in one party, but here in 1933/34, females predominated (51.6%).
In Bol’shezemel’sk tundra, at the beginning of the winter of 1956/57,
the take of males was 56.4%, but by the middle of winter, there was
no difference. On the Taimyr peninsula, at the beginning of the winter
of 1931/32, 64% females were caught. Among 2182 arctic foxes ob-
tained on the Commander islands, males constituted 53.8%.

Under natural conditions arctic foxes are usually monogamous,
but individual cases of polygamy are known, it being observed on
the Commander islands each year. This capacity of arctic foxes is
used in breeding the animals on farms where only one male is kept
with 4—5 females; individual males can accomplish 20—25 matings.
Cases of polyandry, observed on the Commander islands, are ex-
tremely rare.

The ratio of young to adult animals in arctic fox populations
changes considerably depending on feeding conditions, time of year,
and on their total number. On the Lena, 62.9% young were caught

*Not in Literature Cited.; probably Lack, 1954, p. 46—Sci. Ed.

243

348

in the good year of 1935/36. In the Kara region in the season of
1956/57, a quite abundant one, during the time of peak migration
more than 2/3 of the females were young®; in the second half of
winter, the latter constituted only 1/3 of the animals taken. In Yamal,
the prevailing majority of individuals in the populations (in three
harvest seasons from 1955/56 to 1957/58 inclusive; 475 individu-
als—94.7%), were determined as juveniles*. The ratio of adults to
young animals constituted 1: 18.9; this explains the selection of
harvesters in direct ratio to the young animals (Smirnov, 1959,
1959a). Such abundance of young animals over a duration of 3
years otherwise seems improbable.

Arctic foxes reproduce well for 1—2 years in succession, atter
which they “rest” for 1—2 years. This periodicity observed
repeatedly in the 1930’s in the Kola fox sovkhoz was explained by
“physiological exhaustion” of the female organism in which the
first symptoms of overwork were revealed in the third year after
the first two; the older the female, the lower the probability of its
participation in reproduction at this age. Arctic foxes of late litters
(in captivity) do not always reproduce the next year.

By means of the artificial insemination of silver-black foxes
with arctic fox sperm, the mating of female red fox with blue arctic
fox, and blue arctic fox female with the silver-black and white-
muzzled foxes, hybrids of arctic fox and red fox were obtained.
These hybrids had the intermediate characteristics of both species
but all of them were shown to be sterile.

Growth, development, and molt. White arctic foxes are born
covered with dark smoky-brown short fur, while blue arctic foxes—
with dark-brown, almost chestnut. As-in young red foxes, newborn
arctic foxes are born blind, without teeth, and with ear pinnae
closed. They are smaller than [newborn] red foxes and weigh from
60 to 85 g; their body length is 11—13 cm; tail, 4.2-6 cm. Soles of
paws are bare, but already by the fourth day they begin to develop
fur.

Growth in the animals proceeds more intensively than in young
red foxes. Eyes open on the 9th—18th day, simultaneously with the
opening of the external auditory meatus. Eye color darkens over
time. In the blue arctic fox, the eyes are bluish-gray at an early

Uterine determination of age (Chirkova et al., 1959).
*The Russian word sevoletki literally means “this summer’s”—Sci. Ed.

349

age; they turn yellow in approximately the 7th month. On the 12th
day, the irruption of the incisors can be noted, and in some pups,
the carnassial teeth also. On the 15th day all the incisors on both
jaws have irrupted. Canines appear first in the lower, and then in
the upper jaw. On the 18th day, the premolars are cut, first in the
lower jaw. All milk teeth have appeared by the 27—28th day. Re-
placement of [deciduous] teeth by the permanent ones takes place
in wild arctic foxes in the end of summer and the beginning of
autumn in the cross and blue fox types (see below) (Zhitkov, 1913;
Kozlova, 1933).

In one-year-old animals on Yamal in winter, the teeth are snow
white and bright without any sign of wear. In contrast to the red
fox, tooth wear with age begins with the upper incisors, proceeding
to the first and second molars of the lower jaw. Earlier than in red
fox, the fourth upper premolar is worn down and, later than in this
species [red fox], the first and second molars of the lower jaw [are
worn down]. Age determination by tooth wear was shown to be
possible by also taking into consideration the changes in the propor-
tions and configuration of the skull (Grigor’ev and Popov, 1952).
Age determination according to the degree of closure of the central
canine cavity is also possible. In this case, only the first two years
age groups can be clearly differentiated (Smirnov, 1959, 1959а).

With age, the skull undergoes changes. In the young continental
arctic foxes still with milk teeth, the posterior region of the skull is
higher, and the nasal region is shorter and wider than in adults.
Orbits smaller. Zygomatic arches thinner and narrower, bullae osseae
more swollen and rounded, hard palate wider, sagittal crest absent
or weakly developed. Later, the skull broadens in the area of zy-
goma and frontal region, which is abruptly elevated. The postorbital
processes are large and the postorbital constriction is well defined.
Muzzle elongates, sagittal crest is greatly enlarged, especially no-
ticeable in Commander arctic foxes (Ognev, 1926).

At the age of 5—7 years, all teeth in Commander arctic foxes
are worn down. Duration of life of arctic foxes assigned to island
husbandry and sovkhozes for breeding service is 6-10 years. А
case is known of an arctic fox living in a zoo for 20 years, when
the animal died in a state of complete senility (Paramonov, 1929;
Boitsov, 1937; Il’ina, 1949-50). Under commercial conditions, arctic
foxes very rarely survive to old age. In the forest-tundra on the
Yenisei, a very old arctic fox was caught in which only the crooked

244

350

remains of completely obliterated teeth were left; the pelage was
short—about 2 cm in height, tail thin, but the animal itself looked
very large (A.F. Chirkova). On Yamal, in the winter of 1958/59,
when the number of young was small, only 5.5% of the arctic foxes
caught were older than 3 years of age (Smirnov, 1960).

Arctic foxes grow quickly, especially at first. Thus, the average
daily weight gain on farms decreases from 13% of body weight in
the first days to 2.5% at the age of 2 months. Increase in weight
depends also on food composition. On Kil’din Island husbandry on
a mixed ration gave a daily weight increase of 20—40 g from days
42 to 90, an average of 28 g; those kept on fish ration, gave an
average daily weight increase of 41 g and those on seal meat—54
g. Increase in length changed also. In farms, the average weight of
pups at the age of one month was equal to 600—650 g, at two
months—1.7—1.8 kg, at three—2.5 kg, at four—3.8 kg, at five—4.5 kg
and at six months—S kg. At 6-7 months, the young arctic foxes
have almost attained adult size. From the age of 7 months to 2.5
years the weight of arctic foxes on farms increases on average
from 5.6 to 7.1 kg in males, and from 4.9 to 6.2 kg in females.

The change in the weight of the pups of the early and late
litters is greatly different: on a Kola fox sovkhoz, this difference
reached 1.5 kg at the age of 4 months. As compared to the red fox,
the growth constant of the arctic fox before sexual maturity is
larger, and in the sexually mature animals, smaller. Growth of the

“arctic fox is more quickly completed than in red foxes (Boitsov,

1937; Tavrovsku, 1946; Il’ina, 1952).

In nature, young arctic foxes had the following dimensions: on
Yamal, July 2, weight 1250 g, with body length 40 cm and tail 16
cm; August 24 on the Khatanga, body length 51 cm and tail 21 cm;
in October, on the Kolyma, a young blue fox (see below) weighed
3 kg with body length 54 cm and tail 32 cm; the latter animal had
already attained adult dimensions (Zhitkov, 1913; Tsetsevinskii, 1940;
A.F. Chirkova).

Seasonal variations are well manifested. Body weight of arctic
foxes is lowest in summer and highest in December. In connection
with rut and mating, weight drops greatly in the second half of
winter, especially in males. On the Commander islands, it decreased
by 15-20% as compared to that in December—January (Boitsov,
1937; П?’1па, 1940, 1950).

In the white arctic fox, the following age and seasonal stages
of the pelage are differentiated (Paromonov, 1929; Boitsov, 1937;
Guber, 1939; Rakhmanin, 1948; М.М. Spitsyn). Slepushonka* [mole-
vole] or kopanets* [digger] has a delicate dark-brown, almost black
fur, with very sparse, separately protruding guard hairs. Skin length
30-35 cm (without tail); age up to 1—2 months. In the nornik*
[burrow dweller], the quantity of guard hairs is somewhat greater
and the pelage turns browner; the back, shoulders, head and tail
darken. Skin length without tail 40—45 cm. It still lives in the burrow;
age 2—4 months. Krestovatik* [cross fox]—the arctic fox in sum-
mer coat, distinguished by brightening of the pelage on the belly and
along the sides, acquiring a grayish straw-colored tone. The brown
stripe along the back and on the shoulders is clearly marked, in the
form of a cross, and guard hairs are considerably better developed
than in the preceding stages. Skin length of the young cross fox 45—
60 cm, of tail—30 cm. Sinyak* or chayachnik* [blue fox] has a
shorter pelage; following upon the appearance of a large amount of
white hairs, it becomes lighter and grayer, taking on a bluish, leaden
tone. Skin length without tait—54—68 cm, tail length—32 cm. Guard
hairs up to 40 mm in length. Age, 6-8 months. Arctic foxes are
found in this stage from the end of September to the middle of
October. Replacement of the milk teeth by the permanent ones
takes place in the krestovatik and sinyak stages. In the nedopesok*
[premature], fur is grayish-white, with scattered brown guard hairs
concentrated on the upper part of the body. Underfur azure-blue.
The animal is found in this pelage from the middle to the end of
October. Complete, full-grown or doshlyi* [finished| this is the
arctic fox in its winter white fur; some animals only retain a few
“bead” guard hairs, and light bluish color at the base of the hairs.
Wool is dense and fluffy. A few arctic foxes attain the full winter
pelage by the end of November, but the majority in December-
January.

In full winter pelage, the so-called standard arctic fox is divided
into 3 sorts depending upon the degree of maturity, whiteness,
length and evenness of guard hair distribution, and the thickness of
the entire pelage. Not all arctic foxes reach full fur development
each year. The greater part of the skins prepared during the course

*Translator’s remark: These local names (on this and the following pages) are
transliterated, and the literal English meanings given in brackets.

B52

of a winter are of the second and third sorts. In first year arctic
foxes, full development of fur cover is rarely attained. The most
high quality furs are supplied by 3—4 summer [2—3-year-old] males

245 and 2—3 summer [1—2-year-old] females. Emaciated arctic foxes,
and individuals, often frequenting burrows, or living within thickets
of stunted tal’nik [purple osier] are distinguished by dull and frayed
fur.

At the end of winter, when the still white and fluffy fur of the
arctic fox begins to weaken and “leak”, the animal enters the
veshnyak [spring] stage. Later, usually in April, during the forma-
tion of the summer hairs and the darkening of the flesh side of the
skin, the almost guard-hair-less skin is called “gagara” [loon].
After shedding of the winter hairs, the adult arctic fox again enters
the summer stage—krestovatik, which differs from the young ani-
mal by its firmer pelage; length of guard hairs is about 1.5 cm. This
completes the cycle of seasonal pelage change. Throughout the
course of their lives, the adult white arctic foxes annually go through
these described stages of pelage replacement, except for
slepushonka and поник.

The pelage of the blue arctic fox changes with age and to a
lesser extent, with the seasons. From the almost black or dark
brown (slepushonka), the fur of the adult arctic fox gradually
lightens, and acquires a dark violet to light-beige color. With age,
it is mainly the tone of the hair color that changes. At the end of
August on the Commander islands, the majority of arctic foxes
already have a normal “blue” colored winter pelage. Some fur
categories are differentiated—gray or brown tones on Bering Island
and chocolate on Mednyi Island. Hair softness also varies. Arctic
fox fur becomes completely “vykhodnyi” [grown out] in the sec-
ond half of December—January. Average length of contour hairs in
the 1.5—2-month-old arctic foxes is about 27 mm, on winter skins,
more than 82 mm; in winter, hair thickness also increases. As a
result of the warm and humid climate on the Commander islands
and especially in connection with the delayed arrival of cold, the fur
of those arctic foxes is distinguished by coarser and shorter guard
hairs and often by matted underfur (Tret’yakov, 1871; Barabash-
Nikiforov, 1937; Boitsov, 1937; Il’ina, 1949, 1950).

In adult arctic foxes molt takes place twice a year, in spring
and autumn. On farms, arctic foxes have a full winter fur cover

246 from December to February. Spring molt proceeds from the end of

353

Fig. 90. Blue fox in summer fur. Mednyi Island. Photograph by
N.N. Kartashev.

February to June. Summer hairs are initiated in April and grow until
the middle of August. Autumn molt occurs in August-September,
while the growth and maturation of the winter pelage takes place
in October-December. Guard hairs in winter are twice as long as
in summer (an average of 45.8 mm, and underfur 37.9 mm), and
the hair thickness 1$ also greater. Underfur quantity constitutes 97%
(Eremeeva, 1956; Kaletina, 1956; Kaletina et al., 1957). In winter
there occurs an abundant deposit of subcutaneous fat. Sweat glands
at this time are weakly developed. In summer, the fat layer is
reduced and the size of the sweat glands increases.

The beginning of spring molt coincides with the start of estrus.
In the first instance, pregnant females molt, their skins losing winter
fur just after parturition. Males, barren females and yearlings molt
later, and the last are old and diseased animals (A. Romanov, 1941
and others). Holding arctic foxes at higher temperatures (18 + 20°C)
and lower humidity (42 + 8%) speeded up the time of spring molt
by 8 weeks (ша, 1926).

354

In the wild, the time of pelage replacement of arctic foxes
varies greatly depending upon meteorological conditions, the abun-
dance of food and nutritional state of the animals, their age, sex and
the physiological condition of different individuals. Spring molt
begins earlier and occurs more rapidly in years with good food
conditions. In the southern tundra zone, spring molt begins 1.5—2
weeks earlier and ends 2—3 weeks earlier compared to the northern
coast and the Arctic Ocean islands (Rakhmanin, 1959). Molt of
arctic foxes in February was observed in different years on the
Kola peninsula, Yamal and even on Belyi Island. More often, the
onset of molt is observed in March, and on Taimyr and Novaya
Zemlya, even in April. In various populations, it extends for 4 or
more months and in the first 2 months is usually not obvious. Shed-
ding of winter fur ends in the majority of tundras in June or July,
but significant individual deviations are observed. On Klimets Island
(Lake Onega) in the middle of June, a half-white arctic fox was
killed (Sluchevskii, 1886). Individual unmolted arctic foxes are en-
countered even in August. On Novaya Zemlya in the beginning of
May and on Yamal at the end of May, some individuals were found
in winter pelage, and on the New Siberian islands, arctic foxes
were white even at the beginning of July (Krivosheya, 1884;
Paramonov, 1929; Dubrovskii, 1937, 1940; Tyulin, 1938; A. Romanov,
1941; Rakhmanin, 1959). In Greenland (Seton-Thompson, 1909—
1925), arctic foxes are met with which, as in hares, are not able to
change the winter pelage during the short summer and remain
white throughout the year.

Spring molt starts with the shedding of the guard hairs, then
underfur which is, however, more quickly replaced than guard hairs.
Replacement proceeds from the end of the muzzle, rump and tail
base, spreading along the back in the shoulder region, on the limbs
and later—on the side and belly. Winter hairs are retained longer on
the tail. During the formation of the summer hair cover, darkening
of the flesh side of the skin begins around the eyes, at the base of
the tail, on the rump, forefeet and then spreads to the thighs and back.

Autumn replacement of hair cover proceeds in the wild from
September to December. Better fur occurs in January—February.
Time of autumn molt is delayed in the south for 2—3 weeks in
comparison to northern regions (Rakhmanin, 1948, 1959). Thus,
earlier maturation of fur was observed on Belyi Island (about
73° N. Lat.), where at the end of September 1930, arctic foxes

247

355

wandering over the snowless land were sharply distinguished by
their already whitened skins. On the eastern coast of Severnyi
Island of Novaya Zemlya (Sedova gulf, 75° N. Lat., Ledyanaya
Gavan’), during observations on 12 September and October 1936,
arctic foxes in white coats were also encountered (Rundan, 1936;
A.F. Chirkova). Fur maturation proceeds asynchronously in arctic
foxes of various [fur] types. In Kara region, in December 1956,
among arctic foxes of the Pechora type caught, 33.5% of the skins
were of sort 1 [see above], and among arctic foxes of Obdorsk
type, coming from the higher latitude of Yamal—79.7% (Chirkova
et al., 1959). On the Commander islands, maturation of fur at
different times by habitats (“ukhozh”’)* is well known; animals
from rivers in montane barrens have better coats. From the second
quarter of this century, when the climate of the islands warmed
noticeably, autumn pelage appeared later. Earlier, harvest began
here in November—December, when the fur of the majority of arc-
tic foxes was already mature; at the present time, for gathering
better skins, arctic foxes are killed at the end of January and in
February (Il’ina, 1949—1950).

Falling temperature and increased humidity, especially in Octo-
ber, have a favorable effect. Humidity influences increase in guard
hair length and fluffiness. Nutritional status, sex, age, and physi-
ological condition of the individuals affect the time of autumn molt.
It was noted that the more abundant the food in autumn and begin-
ning of winter (but not in summer), the longer the delay in maturation
of fur. Fur matures earlier in males, and later in barren females.
The young of late litters develop full coat later than adults. Molt of
old and diseased individuals is delayed.

Full pelage growth takes place at different times in different
years. On Yamal in the winters of 1923/24 and 1924/25, when the
numbers of arctic fox was low, sinyak [see above] constituted 1.2—
2.0%, 1.е., winter fur of arctic foxes developed comparatively early,
but in 1925/26, when arctic foxes were abundant, this group con-
stituted 20%, nedopesok constituted 18%, and top quality fur was
only 30%; 1.е., growth of winter fur was greatly delayed. In the
winter of 1927/28, when arctic foxes were few, nedopesok con-
stituted 8%, and top quality furs were 45% (М.М. Spitsyn); 1.е.,
growth of winter fur proceeded more intensely. In the lower Lena,

*See previous footnote, p. 346—Sci. Ed.

356

in the winter of 1935/36, when arctic foxes abounded, sinyak were
encountered throughout the winter (in February 0.4%). Skins of
sort 1 in the March collections were 23.2%—development took
place slowly (Tavrovskii, 1939). In the Taimyr National Region in
the winter of 1945/46, in all, 5.3% sort 1 skins were processed
while in the winter of 1936/37—30.6% (N.N. Kuznetsov). On
Novaya Zemlya, individual arctic foxes with dark ears and tail tips
are sometimes seen throughout the winter. Arctic foxes with in-
completely developed winter fur in any case begin spring molt also
in February and March (Tsetsevinskii, 1940). Therefore, in years of
comparatively good food conditions for the young, in years of a low
number of animals, the development of the winter coat is acceler-
ated, while in the absence of food in years of intensive reproduction
of arctic foxes, it is delayed.

For Lena-Khatanga territory the following scheme was pro-
posed for seasonal changes in hair cover of the arctic fox (A.
Romanov, 1941).

January—February—all arctic foxes “doshlyi” [finished], skins
sorts I, П and Ш.

March—predominance of first sort skins; in second half of month,
rarely veshnyak [spring] (the first molt stage).

April—“veshnyak” predominates; in second half of month
“gagara’ [loon] appears (beginning of intensive той).

May——“gagara” predominates (intensive molt).

June—all arctic foxes in “gagara” stage (end of molt).

July—all arctic foxes transit to krestovatik [cross] stage.

August—all arctic foxes in krestovatik stage.

September—4restovatik predominates, a few sinyak [blue],
rarely, nedopesok [premature].

October—krestovatik, sinyak, nedopesok; rarely “vykhodnyi”
[grown out].

November—sinyak, nedopesok; vykhodnyi predominates with
skins of lower sorts [II, Ш]; a few first quality.

December—vykhodnyi skins of all sorts, sort II predominates;
nedopesok is rare.

Other evidence of A. Romanov was also taken into consideration on the
development of arctic fox fur in the winters of 1925/26, 1926/27 and 1934/35, with
different environmental conditions.

248

957

The topography of hair cover replacement in autumn differs
from that in spring. Molt also begins with the head, which turns
gray on the sides, but subsequently, whitening takes place on the
groin and on the sides of the body, and later on the back and tail.
On the back, individual dark hairs are retained for a long time.
Sometimes, whole areas of the fur on the nape remain unmolted.
During the time of the autumn molt the flesh side of the skin in
white arctic foxes remains light, but in the blue, it darkens; at first,
the flesh side grows thick, but later becomes thin.

Among the pelage variants of white arctic foxes, cases of
partial chromism in the form of sharply delimited spots of foxy color
are noted. In the Bol’shezemel’sk tundra, white arctic foxes were
caught with spots of reddish fur on the side or on the posterior part
of the back, from 2—3 to 10—15 cm and larger. Spotted individuals
were observed in the years with an abundance of arctic foxes. In
January 1947, in Nizhnii Kolymsk region, an arctic fox was caught
with a bright-reddish spot 6.3 x 19 cm in diameter.

Enemies, diseases, parasites, mortality, and competitors.
Enemies of the arctic fox are few. Among them are the wolverine
and wolf, which are found in tundra in some numbers. The polar
bear rarely attacks the arctic fox, but in the years of famine, it may
harass the fox to death. The main enemies are sled and herders’
dogs. Among birds, the golden and white-tailed eagles, the large
species of gulls (glaucous and others) and skuas, raven and even
crow are dangerous (to young), and on the Solovets islands, also
goshawk. More often, the white polar [snowy] owl annoys the
arctic fox, being especially numerous in the years of intensive re-
production of the animal; in the taiga zone—eagle owls. Owls attack
young arctic foxes during their dispersal from burrows and also kill
those weakened by disease or migration. Owls are periodically very
abundant in the tundra, Thus, in Bol’shezemel’sk tundra 600 birds
were trapped in spring, summer and autumn 1955 in an area of
about 400 km? (Yu.V. Rybalkin).

In years of insufficient food with an abundance of animals and
intensive migration, epizootics are common in the migrants (Chirkova,
1951, 1955; Sokolov and Cha, 1957; Meteleva and Rubanchik, 1959;
Chirkova et al., 1959). Native inhabitants have long been acquainted
with the disease called here dikovan’, or frenzy [rabies], “plague”.

Losing their sense of fear, furious arctic foxes run into villages,
enter dwellings, attack dogs, reindeer and humans. Animals bitten

249

358

by arctic foxes become rabid, but the disease is extremely rare
among people. Scars and gashes on the muzzle and other body
parts of the arctic fox indicate that the animals often bite each
other, and this influences the rapid spread of the disease. Usually
many other species are involved in a rabies epizootic: red foxes,
wolves, wolverines, ermines, weasels, sled and hunting dogs, hoofed
animals and even birds—owls, gulis and others which feed on lem-
mings. The disease among arctic foxes is usually preceded by the
massive death of lemmings.

The bodies of dead arctic foxes are not often observed in the
winter period, and only after the disappearance of snow is the size
of the epizootic revealed. Thus, in the spring of 1933, in the middle
of Yamal, the remains of 16 arctic foxes which had died in autumn
and winter were found along an extent of 20 km’. In the same
year, in the Se-Yaga region, 130 dead bodies of arctic foxes were
discovered in an area of 3000 km’; the majority were in winter fur
and high nutritional status. In the Pechoro-Ilychsk preserve, in the
second half of winter after the massive migrations of arctic foxes
in 1929/30 and 1935/36, disease was observed with the death by
spring of nearly all immigrants. In the next season, disease was
observed among dogs similar to that of arctic foxes and much
mortality among them (Teplov, 1960; A.F. Chirkova).

Large epizootics among arctic foxes took place, from the end
of 1946 and in 1947 in almost all expanses of tundra in the USSR
after the disappearance of lemmings (following unfavorable meteo-
rological conditions in 1946 and 1947). By the spring of 1948,
surviving arctic foxes were scarce everywhere, reproduction pro-
ceeded slowly, and their numbers sharply decreased by the season
of 1948/49 in all tundras of the Soviet Union, except for the Far
East where their reduction in number had occurred in 1947
(Chirkova, 1955).

At the beginning of the winter of 1953/54, in northern Yakutiya,
soon after an outbreak of lemmings, disease appeared among arctic
foxes. The diseased or dead constituted about 15% of those ob-
tained, and all were well nourished—the arctic fox fed on the dead
lemmings. The disease followed a swift course, and was transmit-
ted to dogs, wolves and reindeer. Later, starvation of arctic foxes,
migration and sharp suppression of reproduction were observed in
1954; about 1/3 of the females were left unmated and more than
half of them remained barren (Tavrovskii, 1958). Only among

359

arctic foxes of the Commander islands were epizootics not ob-
served. The number of animals is always sharply lowered as a
result of epizootics, and the intensity of reproduction of the surviv-
ing animals sharply decreases.

The study of rabies—a neuroviral disease of arctic foxes—was
promoted in the extreme northeast of Bol’shezemel’sk tundra from
1954 to 1962 (Kantorovich, 1956, 1957, 1963). It was established
that outbreaks of the epizootic among arctic foxes took place in
1947/48, 1951/52, 1954/55 and 1955/56, with moderate disease in-
cidence (up to 10 cases) in 1949/50, 1952/53, and 1958/59 and
1961/62, with simultaneous cases among dogs and individual occur-
rences among wolves, red foxes and reindeer. The disease was
observed exclusively in the cold months of the year from Novem-
ber to March. Examination of the brains of 1596 healthy arctic
foxes over the course of 6 summers revealed that in general 30%
were carriers; in years of massive outbreaks or moderate inci-
dence—30—75%, and in the absence of the disease—3—6%. The virus
was also isolated from red foxes. In healthy wolves, ermine, mouse-
like rodents (Siberian lemmings, voles), owls and dogs, the virus
was not detected, and therefore the arctic foxes were considered
the permanent carriers of rabies infection in the far north. The virus
was isolated from young arctic foxes, and from males twice as
often as from adults, or from females. The virus was also isolated
in the parotoid and submaxillary glands in the years of outbreaks in
20-30% of cases, but outside the disease period, in low numbers of
arctic foxes, and in the absence of migration—only in individual
cases.

Rabies in arctic foxes was also studied in the tundras of the
Nizhne-Kolymsk region during the epizootic of 1958/59. It was
shown that in intramuscular infection of 6 arctic foxes from arctic
foxes, red foxes and dogs spontaneously ill with rabies, the incuba-
tion period ranged from 10 to 18 days, after which all arctic foxes
became sick and died within 2-3 days. At that time hydrophobia
disappeared. All animals died in a condition of outstanding good
nutrition. Pathological changes in the diseased animals were re-
vealed only in the form of hyperemia in the brain. Babesh-Negri
bodies were not detected in the diseased arctic foxes examined.
The retention of the active virus in the frozen brain was confirmed
after 202 days. Not a single case of disease among people was
established, even after being bitten by rabid animals. However, the

250

360

question of virus pathogenicity to humans through various routes
and dosages was not studied (Strogov, 1961). The disease is char-
acterized as a viral arctic encephalitis of animals (Stroganov, 1964).

Under natural conditions, the arctic fox is susceptible to
leptospirosis (infectious zheltukha), especially at a young age, but
in a mild form. The role of agent of leptospirosis in arctic foxes, red
foxes and other animals is served by Leptospira icteroanaemiae,
and in addition, this disease in both species is induced by L.
icterohaemorrhagiae (causative agent of Vasil’ev-Wel’s disease
in humans). Animals and humans are both susceptible to leptospirosis
(Vyshelessku, 1954).

Outbreaks of epizootics among arctic foxes was observed in
Bol’shezemel’sk tundra in 1909/10, 1926/27, 1929, 1932/33, 1937/
38, 1943, 1947/48, 1955/56 and 1957/58; on Novaya Zemlya, in
1891, and also in 1927 and 1932/33; on Yamal in 1923, 1928, 1930,
1932/33, 1934/35, 1935/36, 1947/48, and 1957/58; in northern
Krasnoyarsk territory in 1915/16, 1920, 1923/24, 1926/27, 1935/36
and 1947/48; in Lena-Khatanga territory in 1926/27; in northern
Yakutiya in 1944, 1946/47, 1950/51, 1953/54 (from the Yana to the
Kolyma) and in 1947/48 (Olenek-Lena); on the Anadyr in the be-
ginning of the 1880’s, in 1890, 1903/04, and on the coast in 1944/
45.

On fur farms and nurseries [sic], arctic foxes suffer from
paratyphoid which carries off to 50% of the young, especially of the
late litters; this disease has been known in the USSR since 1933
and is caused by the bacterium Salmonella cholerae suis and 5.
typhi murium. There are references to the presence of canine
distemper, Aujeszky’s disease, pneumonia and tuberculosis and rarely
anthrax among arctic foxes. In years of outbreaks of the latter
among reindeer, much mortality was also observed among arctic
foxes (Lyubashenko, 1941; Vyshelesskii, 1954; Sludskii, 1954;
Lyubashenko et al., 1957; Pankov et al., 1957; N.N. Spitsyn).

Skin diseases of arctic foxes in the wild have not been studied.
In years of this epizootic disease, arctic foxes appear with incom-
pletely developed coats, with bald patches and other damage
(Rutilevskii, 1939; Rakhmanin, 1948; Chirkova, 1955 and others). In
the European north, in 1945/46 and 1946/47, bald patches and
“stunted” pelage were observed in arctic foxes. At the mouth of
the Pechora, in February, 1956, at a time of epizootic, an arctic fox
with scabs and almost bald was caught. In tundras of the Far East,

361

in 1944/45, among a great number of sick arctic foxes, up to 70%
“spoiled” coats were encountered.

In two arctic foxes from Yamal in 1957 and 1959, 5 species of
external parasites were found: the gamazid mites Laelaps lemmi
and Hirsutionyssus isabellinus, the fleas Ceratophyllus penicilliger
and Leptopsylla ostsibirica and the biting louse of the snowy owl,
Strigiphyllus ceblebrachys (Luzhkov, 1963).

In arctic foxes of the Commander islands, scabies was ob-
served in some cases, caused by Sacoptes scabiei (0.4%). On
Bering Island, the mange mite (Demodex canis) was observed—a
serious, usually chronic skin disease (7% animals infected), which
was widespread among arctic foxes in the 80’s of the past century
and which was, probably, transmitted from dogs. On the Com-
mander islands, the ear tick Otodectes cynotis and the biting louse
Trichodectes canis were found on arctic foxes; sucking lice were
observed [pediculosis] (1.6%) and they were struck with ringworm
[trichophytosis] and moist ringworm (0.7%) infections. In 1935/36,
of 61 arctic foxes examined, 27 were observed with these diseases
and in 34 arctic foxes the biting louse and sucking louse were
found.

Infection with endoparasites among arctic foxes is almost 100%
(Isaichikov, 1931; Dubrovskii, 1937, 1940; Kirpichnikov, 1937;
Kolmakov, 1937; L. Popov, 1939; Rutilevskii, 1939; Shastin, 1939;
Afanas’ev 1941; Petrov, 1941, 1958; A. Romanov, 1941; Pereleshin,
1943; Il’ina, 1949, 1950; Dubnitskii, 1953, 1957; Chirkova et al.,
1958, 1959; Britov, 1959; Petrov and Kosupko, 1959; Ovsyukova,
1961; Luzhkov, 1963). In the USSR, more than 30°’ helminth spe-
cies were found in arctic foxes, of Nematoda (11 species), Cestoda
(10 species) and Trematoda (7 species) and two species of
Acanthocephala (found in arctic foxes on fur farms).

The nematode Тохабсат5 leonina is found in the intestine and
stomach of arctic foxes nearly everywhere, and in the majority of
animals; of 59 arctic foxes from the neighborhood of Kara, 84.8%
were infected, on Yamal, of 118 animals in a year of depressed
numbers of arctic foxes—92.4%. On the Commander islands, the
fluke Maritrema sp. (80—-100% infection) and the cestode
Mesocestoides lineatus (88% on Bering Island) are found; each

’There are 55 species of parasites known; one species of Protozoa, 44
species of parasitic worms and 10 species of external parasites (Luzhkov, 1963).

—

362

individual contained some thousands of both species. The nematode
Uncinaria stenocephala is very often met with, and infects young
animals more intensively. The nematodes Spirocerca lupi and S.
arctica invade the stomach (goose egg-size tumors); the latter
induce extreme emaciation and susceptibility to rabies. Trichinosis
is known in 34% of some populations.

A great danger presented by arctic foxes is their infection with
the cestode Alveococcus multilocularis, found in the adult stage in
the intestine of 50.8% (among 59) of arctic foxes in the winter of
1956/57 in the northeastern Bol’shezemel’sk tundra; in some ani-
mals up to 200 thousand specimens were found. On Yamal, the
infection rate in 1957—1959 reached 73.7% (in 118 arctic foxes,
from 11 to 74 thousand specimens were found in each animal). In
the Anadyr and Jul’tin tundras (Chukotka), the total infection was
62% (among 35 arctic foxes) with intensive invasion by hundreds
and thousands of specimens (Ovsyukova, 1961). Apparently, it is
this form of echinococcus that is found in arctic foxes on Kil’din
Island, the Commander islands, and in continental arctic foxes of
Yakutiya. This parasite, when present as large infection in arctic
foxes, induces extreme emaciation-and paralysis. On Yamal, arctic
foxes are also heavily infected with the tapeworm Taenia
polyacantha (in 86.4% of animals) and 7. crassiceps (in 46.6%).
Trichinella spiralis, which induces a rise of temperature and pa-
ralysis of respiration resulting in death or chronic illness, was
found in arctic foxes of the Commander islands and in some other
regions of the Far East (Petrov, 1941).

Species of massive infection—echinococcus and the majority of
taeniae—are transmitted through small rodents. The burrows of arctic
foxes are the foci of toxascaridosis. Echinococcus as well as the
trematodes Opistorchis felineus and Metagonimus jokogawai,
the cestodes Diphyllobothrium latum, D. erinacei and D. caninum,
and the nematodes Trichinella spiralis and Toxascaris leonina
are pathogenic to humans also.

The majority of helminth species of arctic foxes are found in
the gastrointestinal tract, mainly in the small intestine (in 100% on
Yamal); one trematode species—in the bile ducts of the liver, 1
nematode—in the urinary bladder, 2 nematodes—in the bronchi, tra-
chea and nasal cavity, and | nematode—in the stomach. The larval
stage of one species of cestode lives in subcutaneous cellular tis-
sue, and one trichinellan—in the muscles. Coccidians were found in

252

363

the liver and intestinal tract of the arctic fox. Each arctic fox
(Bol’shezemel’sk tundra, Yamal) is infected by up to 4 species of
endoparasites. The degree of parasitic infection on the Commander
islands is related to the nutritional condition of the animals
(Afanas’ev, 1941). In almost all arctic foxes, enteritis is noted as
a result of the effect of helminths; in the presence of heavy infec-
tion with alveococci, there is subacute necrotic enteritis. Helminthiasis
reduces fertility and induces the death of animals. Helminths of
arctic foxes are shared in common with red foxes, wolves (4 spe-
cies on Yamal) and dogs. Infections of arctic foxes in the northern
regions are less than in the south (L. Popov, 1939; Rutilevsk1i, 1939;
Chirkova et al., 1959).

Death of arctic foxes due to basic hardship frequently takes
place in the cold period of the year. During ice-glazed periods, the
arctic fox’s claws become broken and it cannot obtain food in the
littoral zone and the tundra. Arctic foxes caught on the thin autumn
ice of the open sea often perish in great numbers. Along the west-
ern coast of Yamal and in Baidaratsk bay, dead arctic foxes are
collected in tens or hundred in some years. Many animals die at the
time of seasonal migrations in the unusual situations of the forest
zone and Arctic seas, whence they come in hundreds in autumn,
and in spring return individually. Arctic foxes often drown crossing
large rivers, for example, the lower Pechora with its numerous
branches. On the Nizhnii Taimyr, of 46 arctic foxes swimming
across, an observer saw three drift out to sea and three die in the
river.

Several facts attest to the great vitality of arctic foxes. On the
lower Lena, an arctic fox was noticed with a badly broken paw;
gangrene had already appeared, and it weighed only 1075 g. On the
Commander islands, a blind female with cataracts in both eyes,
stumbling against rocks while running, brought into the light a litter
and nourished them.

On the Commander islands, arctic foxes died from eating salt
fish. Dead animals were found with pathological changes in the
kidney and with ulceration of the stomach and intestine; cases of
pneumonia and emphysema were noted. Sometimes inexperienced
mothers bite, and tear at, or drag about their young. Deficiency of
milk may incite such mothers. Newborn young die mainly in the
first days after parturition. Mortality among pups up to 10 days old
constituted 22-24%, from 10 to 30 days old—3%, from 1 to 2 months—

364

8%; i.e. the total number of deaths among young at the end of
lactation was 34% (of those born). From 2 to 5 months of age, 3%
of the whelps died, and total mortality through the period of attain-
ment of sexual maturity—about 40%. On the Commander islands
mortality through yearling age of 33% was noted; on farms in
Canada—33—46%. In the Commander island enterprises in the 193075,
mortality among arctic foxes in July—August around an “ukhozh”
[foraging site] varied from 8 to 15%. Mortality among adults con-
stituted about 5% per year.

On Mednyi Island, of arctic foxes that died in the period of
1930—1935, 20% died from injuries and fights, 13% from poisoning
by salted products, about 37% from diseases and 30% from other
causes. On Bering Island, 21% of the arctic foxes taken in 1937/
38 were diseased.

Under natural conditions of the continental tundras, the magni-
tude of mortality, like that of reproduction, varies considerably in
various years as a result of the great fluctuation, above all in the
food supply. Judging by post-parturition placental scars in the uteri
of arctic foxes in northeastern Bol’shezemel’sk tundra in the sum-
mer of 1956 (a year with decreased number of arctic foxes), an
average of 3.1% embryos were resorbed per pregnant female; 50%
of the young females had died by the beginning of harvest. On
Yamal in the summer of 1933, with negligible numbers of lemmings,
about 80% of the young died. Almost a two-fold reduction in the
number of young at the burrows was noted by “Sluzhboi Urozhaya
[Harvest Management]” from May to September in various tundra
regions of the Far North. To control mortality of young arctic foxes
in the wild, local kolkhoz [collectives] and sovkhoz [state farms]
bring food to the burrows in food-deficient years in spring and
summer, as well as in autumn in places of approach and harvest of
the animals (Guber, 1939; Tsetsevinsk1i, 1940; G. Naumov, 1941;
Skrobov, 1958; Chirkova et al., 1959).

During the past decade, a northward shift in the southern limit
of reproduction has been observed. A series of authors (Rozanov
et al., 1932; Skrobov, 1958, 1958a, 1960; Uspenski, 1963) consider
this to be due to the colonization of red foxes in the forest-tundra
zone, and to its deep penetration into the southern zone of tundra.
However, the importance of the red fox as a competitor of the
arctic fox is exaggerated, since the main food of the red fox is gray
voles [Microtus] and of arctic fox—lemmings and marine flotsam.

253

365

Arctic foxes prefer to den in open, elevated parts of the tundra,
while for the red fox it is river floodplains, mainly forested sections.
The number of red foxes in the zone of contact of the ranges of
both species is extremely small; in northern Arkhangel’sk district,
in preparing skins during 28 years (until the season of 1962/63) for
one red fox there were more than 10 arctic foxes and the differ-
ence between the number of both species increased in recent years.
The process of displacement of the limits of reproduction of both
species is governed, one may suppose, by changes in a series of
abiotic and biotic factors as a result of Arctic warming; i.e., the
effect of warmth on the soil and its moisture, on the microclimate
of the burrow, on the duration of snow cover, on the time of the
vegetative growth period, perhaps on the displacement of the limits
of some plant and animal forms, in particular, the distribution of
species of small rodents—lemmings, which are the main food of the
arctic fox and gray voles, preferred by the red fox. Attacks on
young arctic foxes by red foxes were observed, as well as of arctic
on red foxes, in the course of ousting of one or the other species
from burrows and places of activity (Rozanov et al., 1932; Boitsov,
1937; Dubrovskii, 1940; Skrobov, 1958, 1958a).

Direct competitors of the arctic fox for food are the ermine and
weasel, but because of the small number of these species in the
tundras, they do not inflict large losses on the arctic fox. In the
lower Khatanga, where the arctic fox appears in great numbers
there, the ermine disappears. In years of lemming abundance, they
are also eaten by wolf, polar bear, wolverine, wild and domestic
reindeer and even shrews, and thus they compete with the arctic
fox. On Kil’din island, contests with river otters for fish were
observed, and in the littoral zone of the Commander islands—with
swines for marine flotsam. Sometimes, the wolverine uses burrows
of the arctic fox. In its turn, in Yakutiya the arctic fox occupies the
empty burrows of the tarbagan marmot and in the mounds of ground
squirrels.

Everywhere in the tundra, the snowy owl competes with arctic
fox for its main food which is also the small rodent; in years with
many rodents, both the snowy owl and the arctic fox reproduce in
great numbers in the tundra. The same was observed for the rough-
legged hawk [Buteo lagopus], which is less numerous in the tundra.
Small rodents are also eaten by 2—3 species of skuas [Stercorarius},
peregrine, merlin, in some places short-eared owl, white-tailed eagle,

366

goshawk and, in addition, raven, crow, magpie and gulls; gulls and
corvids feed together with the arctic fox in the littoral zone
(Osmolovskaya, 1948).

Population dynamics. Seasonal dynamics in numbers of the
arctic fox in years when its main food is plentiful is well mani-
fested. Minimal population density is observed in spring, and the
maximum at the end of summer—beginning of autumn, just after the
litters abandon the burrows. When food is scarce, the growth of the
population in summer is sometimes so negligible that the number of
arctic foxes in autumn is even less than that in spring.

Annual variations in their numbers are extraordinarily sharp.
Beginning in the middle of the last century, a 3-year repetition of
“productive” years for the arctic fox was often noted, with fluc-
tuations of from 2 to 6 years between years of abundance (Vrangel’,
1841; Brandt, 1856; Middendorf, 1869; Nosilov, 1909; Folitarek and
Chirkova, 1930; Burke, 1936; Dubrovskii, 1937, 1939, 1940;
Kirpichnikov, 1937; Guber, 1939; Rakhmanin, 1948; Chirkova, 1955;
Sdobnikov, 1958 and others). During 35 years at the end of the past
century and the beginning of the current one, the maximum supply
of arctic fox skins at Irbitsk fair was observed each 3—5 years, and
at Yakutsk each 1—5 years (Silant’ev, 1898; Paramonov, 1929).
During 1924—1959, maxima of prepared skins were repeated each
2-4 уеагз*. In separate regions of the north, after the October
Revolution, the interval was 1—4 years (2—3 years on the average).
During the last decade, some stabilizing of fluctuations is observed
in the arctic fox catch. Apparently, this is a result of technical
measures to improve hunting. The average catch of arctic foxes in
this period increased (see p. 376). Fluctuations in the catch and in
preparation of skins correspond, in general, to the dynamics in the
number of the species revealed by the more or less accurate count
of wild animals in nature.

There is great amplitude of variation in the numbers of arctic
foxes in successive years. In the autumn of 1912, in the vicinity of
Russkii Ust’ [Russian Mouth] of the Verkhoyansk region there was
an extraordinary “raid” of arctic fox, “what mosquitos” as ex-
pressed by the natives (Zenzinov, 1914). Following such a year of

58Not counting the war years, when in the USSR as a whole and in Krasnoyarsk
territory, 5 years passed between 2 successive peaks, probably as a result of
special economic conditions.

254

367

abundance, the arctic fox was rarely met with. The number of
animals in census areas varies tenfold in various years. The number
of breeding burrows changes from a small percentage in unfavor-
able years (Table 29) to full occupation of all breeding burrows and
the appearance of pups even in temporary shelters in years of
abundant reproduction of the species. In 1935, in the Kara region—
in northeastern Bol’shezemel’sk tundra, in a year of growing
numbers of the species, 39% of the burrows found were occupied
by large litters and the total number of arctic foxes on the experi-
mental plots increased more than 9 times as compared to the
preceding year, when only 5% of the burrows were occupied and
litters were fewer. In the same region in the winter of 1957/58, as
compared to the preceding winter, the number of arctic foxes de-
creased 3.7 times (N. Dement’ev, 1955; Chirkova et al., 1959). On
the northern shore of Taimyr Lake and along the Nizhnii Taimyr
river, with intensive reproduction of arctic foxes in 1947, about 70%
of the old burrows were occupied; in 1948, with weak reproduction,
out of 18 old burrows examined, a litter was found in only one
(Sdobnikov, 1958).

In the arctic tundra of the Nizhne-Kolymsk region in the sum-
mer of 1950, the greater part of the burrows were inhabited, and
in autumn, the number of arctic foxes was high. In 1951, the num-
ber of arctic foxes sharply decreased and out of 205 burrows, only
11.5% of the burrows were inhabited; at the same time, in the
southern zone of the tundra, where food is more varied, a concen-
tration of arctic fox was noted. In summer of 1952, in the arctic
tundra, out of 103 burrows 2.3% of them were inhabited, while in
the southern :zone—41.2% (of 34 burrows). In summer and autumn
of 1953, with abundant lemmings in the arctic tundra, more than

Table 29. Occupation of burrows by arctic foxes in Turukhansk
territory, around 69—70° N. Lat. (Boitsov, 1937)

Total Of these, in %
burrows
investigated With 3 With signs Uninhabited
pups ———SSSS SSS
Permanent Rare
residence visitation
of adults

7] 2:7 7.0 ты 79:2

256

368

NIN Wy

is Ti Mg 2647 HR в ТП eg “Oy "hy ils "bly Ч о Ба Soy Oe
Tag hy hs $47 «99 505) 3453 245 2057 2659 Об
25 278 240 "Hyg Ho ha ha “Ss Ч “Hy Hp зу. 5566 She “hg

Fig. 91. Tanning of arctic fox skins in the Soviet Union as an index of
fluctuations in the numbers of the species (A.F. Chirkova).

40% of the burrows were occupied by arctic foxes and reproduc-
tion proceeded well (average litters of 5—7 young) (ТаугоузКи, 1958).
As is seen, the numbers of arctic fox do not change synchronously
in the various geographic regions and even separate places, never-
theless, great rises and falls in numbers simultaneously envelop vast
areas of the European and Asiatic tundras eastward to the Lena
[river]. Farther to the east, the rhythm of dynamics does not usually
coincide. Only in occasional years the rise in number is observed
in the area of the whole tundra zone of the Soviet Union, as it was
in 1922/23, 1929/30, and apparently in the 3 years 1935/36—1937/38.
A “crop” of arctic foxes in the winter of 1944/45 was noticed in
the whole North with the exception of the Far East.

_ Even in the last century, the dependence of the numbers of
mainland arctic fox on the abundance of lemmings, as the main
food of arctic fox, was noted. At the present time, it has been
convincingly shown that the massive appearance of lemmings stimu-
lates reproduction in the arctic fox only when it takes place in the
preceding autumn and winter (subnival reproduction of rodents). In
the whole north of the Soviet Union, during the period 1944—1948,
there can be traced a clear dependence of reproduction, migrations,
and development of epizootics, and, as a result, the status of the
numbers of arctic foxes, on the abundance of lemmings in the
preceding autumn and winter, and of these rodents on the charac-
teristics of weather in the course of different seasons of the year
(Fig. 93). The value of these interrelationships and governing rules
allows one to make a prognosis of numbers and population status
of arctic foxes some months before the beginning of the harvest

255

в ей .: Sh
A BGG
LE amen ra

x
x

о
ee

тж
Vd <
ты
i
= My ie

Oo
rss

ES Se Е ZH:

ge We a
ESET ae aan

369

Fig. 92. Variations in relative numbers of the arctic fox in the tundras of
the Soviet Union over 5 years—1944/45—1948/49 (from Chirkova, 1955):
1—low number; 2—number below average; 3—average number; 4—number

above average; 5—high [number].

Temperature HUTT TTT A
Precipitation [IMM ZA MUN | IZ

256

370

}
—
2244

1
>

‘Numbers of arctic

fox: and
© skins as % of
1944/45 season
ra
=

= Tanned arctic fox

lemmings

Е]

<
Ss

.] i(

TUTTO TOT eee
GYGoNGs
ИА NAA NGRU BBN aa AUN

WI aa | AA A LAGI SOOO SSO

Months ГЛУИДЛ1ДУИЕЛТПУИДЛ ГИУИКЛ 1 OY

со
i=)

Moderate | 50

VIM TITI LLL LLL LL LIL LLL LLL LLL LLL LIAL L AAA

VTILLLLLLLLLL LALLA LALA LLL LLL ALLA LA LLL LL

=)
te
(a)
22

Рем | 0

RS
АК
<

Rye
as
ae
SN
ы
§
I

К
ПИШИ ЛИДИИ ИХ ИИ
Years 1943 1944 1945 1946 1947 1948

Fig. 93. Weather condition, reproduction of pestrushek* (lemmings) and
number of arctic fox in the tundras of Yamalo-Nenetsk zone during the
period 1943—1949 (from Chirkova, 1955).
1—numbers of arctic fox; 2—numbers of lemmings; 3—tanned arctic fox
skins; 4—prognosis of the number, based on sum of indexes; S— favorable
weather (precipitation, temperature); 6—moderate [weather]; 7—unfavorable
weather (for lemmings).

season (Dubrovskii and Romanov, 1935; Romanov and Dubrovskii,
1936; A. Romanov, 1941; Chirkova, 1951, 1955).

Migrations have great significance in the dynamics of arctic fox
numbers. As stated above, through this, emigration of huge num-
bers of the animals from the tundra zone and their appearance in
the taiga zone and on the Arctic Ocean, takes place. Latitudinal
migrations of arctic foxes in some regions, for example, in Kanino-
Timan tundra at the time of great increase in their numbers, increases
the total “crop” of arctic foxes, while the local groups of the
animals remain small (Bashkin, 1940, Shibanov, 1951). On the

*Local name—Sci. Ed.

ЕЕ

257

371

Commander islands and the sea coasts of the mainland tundra
in the east and west of the range in the USSR, where the food
supply in the form of marine flotsam is relatively permanent, the
number of arctic fox is more stable and changes less in the various
years.

Exploitation [for fur] also reflects numerical dynamics. Inten-
sive trapping out of arctic foxes in years when they are abundant,
increases the probability of survival among the remaining animals.
Increased take of arctic foxes in the years when they are few in
numbers is not rational, since it decreases the producer group.
Hunting of arctic foxes in the burrows in the second half of winter,
when they are preparing to breed, is particularly harmful. Rational
utilization, taking into consideration the numbers of arctic foxes and
the conditions of their existence, may favorably influence the popu-
lation, increasing its productivity.

Blue foxes, common in regions with mild climate, and extremely
rare in mainland tundra (a tenth and a hundredth part of the per-
centage among white arctic foxes; Paramonov, 1929; Lavrov, 1932;
Zubkov, 1936 and others), are more often observed in years of
abundance of the species. In Canada, blue arctic foxes also occur
in larger quantities in years of massive reproduction of white arctic
foxes (Hewitt, 1921).

In Canada, massive purchases of skins are repeated each 3—5
years, 4.2 years on the average. The catch of arctic foxes in
individual years decreased 8—9 times (Seton-Thompson, 1909—1925;
Hewitt, 1921). Co-occurrence of the year with an abundance of
lemmings was established in both Norway and Canada (Elton, 1931,
1931a). The numerical dynamics of the arctic fox in northern and
regions of the Arctic and sub-Arctic zones of America differ.
Increase in numbers of arctic foxes over large territories in Canada
was noted in 1933/34, 1937/38, 1941/42 and 1945/46, and massive
reproduction of lemmings—in the same or the preceding years (Elton
and Chitty, 1937, 1938, 1939, 1940, 1941, 1942, 1943, 1950).

Stocks of arctic fox in Greenland are subjected to strong annual
fluctuations depending on the arrival of spring, meteorological con-
ditions which are determined by the amount and character of the
passing of huge masses of drifting ice from the Arctic Ocean to the
south along the shores of the island (Vibe, 1959).

258

372

Fig. 94. Track of an arctic fox on damp sand. Khatanga. Photograph by А.Е.
Chirkova.

Field characteristics.» The narrow paws of arctic fox leave
tracks similar in form and dimensions to those of the red fox, but
somewhat more blunt (Fig. 94). On the firm snow of the open
tundra, the arctic fox moves about freely, but in the soft snow of
the forest zone it sinks in. Due to its highly fluffy paws, the con-
tours of winter footprint are somewhat diffuse. The length of the
track of the fore limb is 5—7 cm and of the hind, 5—8 cm. The track
of female is smaller than that of the male. The load on 1 cm? of
the supporting surface of the limbs is 40—60 g.

The tracks of all four paws are sometimes distributed sepa-
rately or in pairs (Fig. 95). The usual movement of the arctic fox
is a slow trot or easy gallop with the stride length from 45—50 cm
to 100 cm or more. When pursued, the arctic fox develops greater
speed. In times of fattening it, like the red fox, very thoroughly

58Middendorf, 1869; Nosilov, 1909; Cherskii, 1919—1920; Formozov, 1929,
1946, 1959; Barabash-Nikiforov, 1937; Dubrovskii, 1937, 1940; Tyulin, 1938;
L. Popov, 1939; Rutilevskii, 1939; Shastin, 1939; A. Romanov, 1941; N.
Dement’ev, 1955; S. Popov, 1957; A.F. Chirkova.

373

Fig. 95. Tracks of a running arctic fox on a snow-covered surface. Khatanga.
Photograph by A.F. Chirkova.

searches all bushes, hummocks and projections above the soil sur-
face. During the hunt, the arctic fox changes its gait, now slowly
sneaks up, sniffing a track or following the prey at a distance, now
creeping on its belly, then quickly glides forward or makes a sudden
jump to overtake the victim. The arctic fox seeks out lemmings
along their runways, senses and hears through the mass of snow,

259)

374

vigorously digging them out even in the compressed snow up to 1
m deep. At the capture site, only traces of blood or the stomach
of the rodent remain. On Belyi Island in 1936, 49 holes 10—15 cm
deep each were dug in the snow by an arctic fox in an area of 100
m’, who made 3 passages in 50 cm reaching the soil itself; for all
of this only one lemming was caught.

In distinction from the red fox, the arctic fox makes broad
trails, especially during mass movements along the seashore and
river banks. On Novaya Zemlya, in years with abundant arctic
foxes, one may, according to Nosilov (1909), travel by horse along
their trails. On ice, the tracks of [polar] bear are often accompa-
nied by the tracks of the arctic fox (Vrangel’, 1841; Kirpichnikov,
1937 and others).

In the second half of winter, double and triple common trails
and sometimes even tracks of four animals running together are
often encountered. Traces of mating and fighting most often may
be seen in the Lena-Khatanga territory in February; at this time,
burrows emptied of snow are found. In the tracks of females,
mucous secretions are met with—signs of estrus. After mating,
arctic foxes lead a secretive way of life, and signs of their activity
are rarely encountered at that time. With the bright sun of spring,
the white arctic fox is difficult to distinguish on the snow. On the
contrary, the blue arctic fox looks almost black.

The breeding burrows are recognized by their well-worn* exits,
the trampled vegetation, the digging and trails of the young arctic
foxes, and the abundance of droppings and the remains of food.
The presence of young in the burrow is betrayed also by the in-
tense odor. Arctic foxes may be revealed by tapping on the soil
surface above the burrow; young arctic foxes reveal their presence
by short barks at such a time, while adults remain quiet. By the
number of active entrances, one can judge the number of young.

At the end of summer, after the dispersal of the litters, and in
autumn at the time of migration of the arctic fox, the river banks
and fast shore ice become dotted with their tracks. Before the
rivers freeze or formation of fast shore ice in the sea, tracks of the
animals are concentrated in a narrow 1-2 т strip along the edge of
the water. With the growth of fast shore ice, the arctic foxes move

*The Russian word obglazhennyc literally means either “picked,” “gnawed,”
or “smoothed,” “worn’—Sci. Ed.

375

with it farther away from the shore. They move out onto the
thinnest ice, and only the sound of the ice cracking makes the arctic
fox instantly jump backwards; during this, it raises its pads like a
cat, fearing to wet them. To the side of the general path, only
tracks of individual arctic foxes are observed, running hurriedly to
a ravine or to the thicket of bushes, in order to search for food.
From the tracks, one may see how the arctic fox can, with a
sudden jump punch a hole in the thin ice to catch a small fish under
it. A dog following an arctic fox onto newly-formed ice often falls
through, while the arctic fox, being lighter in weight, only slips.

The sense organs in the arctic fox are very well developed.
The arctic fox can hear the movement of the small rodents under
snow from some meters; it can react to bait at a distance of 300—
400 m and run precisely in the required direction. It can see willow
ptarmigan far away, entirely invisible on the white snow cover of
the flat tundra. The sense of smell is excellently developed in the
arctic fox; it can scent a capercailzie in the snow at 100 m. When
the animal seeks food, its head hangs down and the nose “traces”
the path. In its absorption in the search, especially when young, the
arctic fox often does not pay any attention to surrounding objects
and approaches a stationary human to within 2-3 т.

The arctic fox does not belong in the number of silent animals,
and gives voice in various situations. The sounds given by it are
quite varied. The usual cry of the arctic fox resembles that of a
-dog’s bark, but more hoarse, deep and husky. An arctic fox hidden
among rocks grumbles “urin-urin”. Caught during migration, sitting
in a sack, the arctic fox gives a cry “ugkhu-ugkhu” when ap-
proached by a human. During the time of rut, females howl, yelp
and squeal. When young arctic foxes disperse from the burrow
prematurely because of lack of food, the agitated parents call them
with a cry similar to that of the swan; this is most often heard at
dawn. Hunters say “The arctic fox coos”; such a cry is consid-
ered to be a forerunner of migration.

On Novaya Zemlya, in the autumn of 1887 and 1888, the bark
of arctic foxes was often heard at the time of migration. While
running it might pause, and, raising its snout, it barked not strongly,
but “complainingly”. Arctic foxes also howl while running in win-
ter on lake ice. Judging from everything, howling in arctic foxes is
related to deficiency of food. (A. Ch.).

260

261

376
Practical Significance

The arctic fox is one of the valuable species of fur-bearing animals.
The USSR occupies first place in the world in the size of its fox
fur trade. In pre-revolutionary times, tens of thousands were ob-
tained in Russia and about 3/4 of the harvested catch was sold in
internal fairs, of which the chief were the Irbitsk, Nizhegorod and
Yakutsk. Part of the skins were exported to China, Manchuria,
Japan and America. There is information on the exportation of
great quantities of arctic fox skins from some regions of Russia. In
the middle of the 18th century, according to Steller, more than 7000
arctic foxes were caught during three wintering periods on Bering
Island. In 1825 and 1826, 140 thousand skins were exported from
Turukhansk territory; from Mangazeya, up to 40 thousand. Indi-
vidual authors refer to this in connection with the decline in the
harvest of arctic fox. The decline in catch of arctic fox was par-
ticularly sharp in the last years of the Tsarist government.

Organized exploitation of arctic fox in the USSR began in 1922
(Paramonov, 1929; Guber, 1939; A.F. Chirkova). During the last 35
years, tanned arctic fox skins in the USSR consisted of an average
of 78 thousand per year, ranging from 35 to 128 thousand. From
1946/47 to 1958/59, the average amount of skins tanned in a year
increased to 81 thousand. This was promoted by the utilization of
new areas, provision of supplemental feeding* of animals, the tak-
ing and holding in captivity of migrating arctic foxes, and also the
rationalization of methods and tools of exploitation.

As regards the price of furs, before the October Revolution,
the arctic fox occupied second to fourth place after squirrel, sable
and red fox, depending on the “harvest” (Nosilov, 1909; Guber,
1929; Chirkova et al., 1959). For the last ten years, the arctic fox
stands in second to third place, providing 15% of the price of furs
tanned in the USSR, while in the tundra zones—up to 90-99%, and
representing the main object of hunting trade for native peoples of
the Far North. The most valuable white fox fur is obtained from the
Yenisei tundras, Yakutiya and Novaya Zemlya. The main regions
of exploitation are the Ob’, Yenisei and northern Yakutiya; during
the last 20—30 years they provided 30, 25 and 23% respectively of

*See footnote, Table 25—Sci. Ed.

377

all of arctic fox production in the USSR. Arctic foxes caught in
northern Arkhangel’sk district (14%) and in the Far East (7%) are fewer.

During the above-mentioned 30—35 years, an average of 5.8
arctic foxes are caught annually in an area of 100 km’ of the
tundras and forest-tundras in the Nenetsk and Yamalo-Nenetsk
national regions, in Krasnoyarsk territory—3.2; in Yakutiya—3.7,
and in the Far East—1.2. At the same time, in the territory of the
kolkhoz [collective] “Krasnyi Oktyabr’ ” [Red October], on the
Kara Sea coast, in the 1950’s, an average of 19 arctic foxes were
caught yearly, and in years with a good “harvest’—p to 30. In the
end of the past and at the beginning of the current century, on the
Commander islands, up to 13 arctic foxes per 10 km? were cap-
tured by means of drives (through the year). Success of the harvest
of arctic foxes differs greatly by year. One hunter taking 10—20
arctic foxes in a season of moderate numbers, may catch up to 100
or more animals in years of abundance. Individual hunters in north-
eastern Yakutiya obtained up to 176 arctic foxes, on Wrangel’
Island—up to 125, and in northeastern Bol’shezemel’sk tundra—
more than 200 arctic foxes per season.

In connection with the sharp variation in the numbers of arctic
foxes, there arose the necessity for an organized service to esti-
mate the “harvest” and to make a prognosis of arctic fox number.
It was organized by the Administration of Glavsevmorput’ [Main
Northern Sea Route] in 1935, when the first general prognosis of
changes for the whole North was given, which proved to be true
for a considerable part of the USSR (Dubrovskii and Romanov,
1935; Romanov and Dubrovskii, 1936; Guber, 1939). The institute
of the hunting industry (VNIO) [All-Union Scientific Research
Institute of Hunting], in the course of a series of years, presented
an analysis of the condition of the populations for the tanning
organizations, and gave a prognosis of the number of arctic foxes
in all sectors of our Far North; they were verified in the over-
whelming majority of cases (Chirkova, 1955).

At the present time, the main methods of arctic fox capture
combine old ways and equipment with new techniques. In some
places, the old method of capture are used; driving with [rein]deer,
barriers on the banks of rivers and the sea, but they are additionally
equipped with new types of deadfalls, food-baits, the newest jaw-
traps and explosive weapons. Wooden deadfall traps are mainly
used in the Taimyr National Region and in Yakutiya, and rarely in

260

262

378

Fig. 96. Slopets, or deadfall, for arctic fox (side wall open) in northeastern
Bol’shezemel’sk tundra (European North). Photograph by Yu.V. Rybalkin.
[See also Fig. 97.]

other tundras (Figs. 96, 97). They represent a narrow corridor with
a bait in a box.* The arctic fox is killed by a log, hung above the
corridor. Deadfalls are located along the seashore, river banks and
in other sites, where the arctic foxes are most frequently encoun-
tered. On the Taimyr peninsula, trap-lines extend from the border
of the forest in the south northward deep into the tundra for 100—
150 km and more. At the present time, improved traps are being
deployed, especially portable traps—irreplaceable equipment for har-
vest in sparsely populated tundras. The steel jaw-traps occupy
second place among the implements of trapping. They serve as the
main trap used for harvesting arctic fox in the western tundras and
in northeastern Siberia.

*The Russian word is strozhka, literally “lodge” or “hut’—Sci. Ed.

3779

Fig. 97. View of deadfall from the front. Photograph by Yu.V. Rybalkin.
[See also Fig. 96.]

A pot drive*, or “tolar” is employed on Yamal and in the
tundras of the European North. In it, from 2—6 tens of hunters on
reindeer teams participate, traveling in a spiral over open, compara-
tively level territory of an area of some tens of square kilometers.
Arctic foxes, fearing to cross the fresh trail of the deersled, gather
in the center of the site around which the harvesters are traveling.
The hunters kill the arctic foxes with guns, or even sticks. In years
of abundance they take up to 150—200 arctic foxes in a circuit. The
method of tolars on foot also exists. Sometimes, the arctic foxes
are driven on to promontories of the seashore or into nets that are
set up (Romanov-Il’inskui, 1958; Skrobov, 1963).

*The Russian phrase is zagon kotlom—Sci. Ed.

380

Fig. 98. Blue arctic foxes arrived for supplemental feeding. Mednyi Island.
December 1953. Photograph by S.V. Marakov.

Arctic foxes are frequently shot from a blind, during solitary
stalks on [rein]deer or horses, with dogs or in an accidental encoun-
ter. Harvest of the arctic fox on the seashore during the time of
autumn migration is permitted only in an organized arrangement for
the sake of keeping the animals in captivity until full pelage. For-
bidden everywhere is klyapts*, digging out animals from burrows,
baited crossbow-traps, and baits poisoned with strychnine, corro-
sive sublimate [mercuric bichloride], nuxvomica or poisonnut,
tormentil and other poisons. The modern method of catching live
arctic foxes by means of food-supplied live traps is employed mainly
in the island enterprises, but has begun to be used in mainland
tundras. In Bol’shezemel’sk tundra, portable food-supplied live traps

*Meaning not clear; perhaps a kind of trap—Sci. Ed.

263

381

of simple construction are used (Boitsov and Pereleshin, 1934;
Freiberg, 1939; Skrobov and Kozhevin, 1955, 1957).

The best methods for catching the arctic fox are snares and
traps based on the employment of lures and baits. Lures for the
arctic fox may be visual, food, scent and mixed. Each object on the
snowy surface of the tundra can be seen from afar and attracts the
arctic fox’s attention, and therefore hunters drive into the snow
pieces of deer-antlers, chocks, set up piles of stones, and pile up
hillocks of earth or snow, etc. Food lures are very varied. However,
fresh carcasses of lemmings, grouse, hares and spoiled eggs are a
continual preference of the tundra population of arctic foxes. Arctic
foxes of the coastal regions prefer fresh fish and fat of marine
animals. Blood of freshly-killed [rein]deer and other animals acts as
a good lure. In order to attract arctic foxes to hunting areas before
the beginning of the harvest, they place baits such as carcasses of
common seals, ringed seals, white whales and other animal and
fishing industry wastes to which arctic foxes are attracted to feed.
This considerably increases the catch.

In previous times, the hunt for arctic foxes continued through
almost the whole year. In the middle of summer the young nornik
were dug out of burrows for nursing. The regular harvest of skins
began at the end of summer, taking in abundance krestovatik
[cross], sinyak [blue] and nedopesok [premature], and the catch
continued to April. In years with an abundance of migrating arctic
foxes, a coastal harvest existed, when 5 to 10 or more animals not
in full pelage were taken per night. In such years, more than 50%
of the harvest consisted of “nevykhodnye’’* arctic foxes. In the
past, arctic fox skins were given as tribute, from each Nenets two
arctic foxes were taken, while from Evenk and Dolganin—four. As
a quantitative unit of exchange, among the Vaddeevsk Samoyed, 20
arctic foxes were equal to one boat.

From the first years of the Soviet state, the catch of
nevykhodnye arctic foxes—nornik [burrow dwellers], krestovatik
[cross], sinyak [blue] and the spring “gagara” [loon] was forbid-
den. A harvest season was established from the middle of
November—beginning of December, to the beginning or end of March
(in various regions).

*Literally “unfinished”, i.e. immature—Sci. Ed.

264

382

An arctic fox industry is conducted on islands surrounded by
unfrozen seas (Boitsov, 1937; Il’ina, 1949—1950 and others). Bering
Island supplied arctic fox to Andreyanov Island (Aleutians) which
belonged to Russia at that time (Khvostov and Davydov, 1810—
1812). At the end of the eighteenth century, G.L. Pribylov [Pribilof]
exported 8,000 skins of blue arctic foxes from Georgii and Pavel
[St. George, St. Paul] islands in two years, and only after that arose
the question of the preservation of these animals there. It was of
great practical importance to organize an intensive arctic fox indus-
try on the Commander islands, where blue arctic foxes occur. In
1741, when Bering visited these islands, blue arctic foxes thrived
there in “incalculable packs”. Steller killed tens of arctic fox with
an axe in a day, and used the skins of the animals to cover the
roofs of the earthen huts. At the end of the 90’s of the nineteenth
century, 1.5 to 2.5 thousand arctic fox skins were taken annually
from the Commander islands. In the beginning of the twentieth
century, as a result of overharvest, drives began to be allowed only
in alternate years, preserves were allotted, counts of arctic foxes
were promoted to determine the number allowed to be killed, and
the permissible period of capture was shortened to 2—3 weeks. As
a result of many years of selection, white arctic foxes, individuals
of which appeared on Bering Island particularly often, were exter-
minated. Only under the Soviet government in the 20’s was winter
supplemental feeding introduced and in the 30’s, in summer as well.
Food-baited live traps began to be used and continuous zootechnical
selection conducted. It is supposed that the arctic fox population on
the Commander islands can be increased to 40 thousand head. In
the 20’s, attempts were made to transfer arctic foxes to the small
coastal islands of the Pacific Ocean and to the Solovets (united in
winter with the mainland by an ice bridge), with the aim of orga-
nizing there an island blue arctic fox industry. All of these ended
unsuccessfully.

Experiments on supplemental feeding of the arctic fox showed
the great importance of this measure not only for islands, but also
in the mainland tundra on several farms in isolated territories. Supple-
mental feeding must be begun during the time the litters are still in
the burrows—at the first signs of the disappearance of natural food
in spring, summer and early autumn, to inhibit the arctic foxes from
migrating, and also in winter in places where animals concentrate
and before the beginning of rut.

383

Fig. 99. Blue arctic fox lying in rocks. Bering Island. October 1958.
Photograph by S.V. Marakov.

Intensive arctic fox farming on islands has been conducted in
North America for the past century, based mainly on wastes of the
marine animal industry (Suvorov, 1927; Boitsov and Pereleshin,
1934; Boitsov, 1937). As already mentioned, the first release of
blue arctic foxes on the Aleutian chain was done by Russians in the
middle of the eighteenth century already, and on the coastal islands
of Alaska in 1835, and American trading voyages for this activity
were undertaken only in 1885. Supplemental feeding of arctic foxes
on the Pribilof islands began in 1896; later other measures were
carried out. Concentrations of arctic foxes here attained very high
densities; up to 500 arctic foxes occurred for each food-baited live
trap.

265

384

Commercial raising of arctic foxes in state animal sovkhoz*
was begun in the USSR in the 30’s. Raising the blue form was
introduced rationally due to their great value. Cage raising of blue
arctic foxes, developed in a series of European countries and in
North America, past through a crisis in the post-war years as a
result of the creation of a mode for short fur. At the end of the
1950’s, some rise in demand was again noted, though public taste
turned to white and light-blue arctic foxes.

The proportion of white arctic fox in the fur export of USSR
reached 7.3% (1933). Before the Great Fatherland War**, blue
arctic fox was priced significantly more expensively than white in
the world market, and the darker, the higher, but in the Leningrad
fur auction (1961), white arctic foxes sold with higher prices than
the blue, and among the latter, the more expensive skins were of
light color.

World production of arctic fox fur (Brase, 1925) in the period
1907—1909 averaged 116.0 thousand skins, of which 105.0 thousand
were white, and in 1923—1924, 204.0 thousand skins, of which 166.0
thousand were white. North America contributes on average about
1/3 of the skins of white arctic fox. In the 20’s and 30’s, from 18
to 72 thousand skins were bought in Canada, an average of 44.9
thousand. In Greenland at the end of the last century, an average
of 1.5 to 2.0 thousand arctic foxes were caught every year. In
Spitsbergen in the 19th century, the arctic fox was exploited espe-
cially intensively. In the 20’s of the current century, only 100 blue
and 250 white arctic foxes were obtained annually. At the present
time, great attention is paid there to the preservation of arctic fox
(Soyrinki, 1958; Bergman, 1961).

Arctic fox meat has an unpleasant odor, but on Yamal many
people ate it (Nosilov, 1909; N.N. Spitsyn). At the present time, the
question of the specific activity of the fat of arctic and red foxes

as a medical dietary treatment against tuberculosis and liver dis-

eases in humans arises (Mironov, 1953).

Arctic fox has a negative significance as a disperser and trans-
mitter of tundra rabies. In years of outbreaks of this disease, many
sled- and herd dogs and reindeer are bitten by arctic foxes, are
infected and die. However, the frequency of disease transmission

*State farms—Sci. Ed.
**W orld War I—Sci. Ed.

266

385

by arctic foxes may be exaggerated due to insufficient study of the
significance of rodents and other animals of the tundra. Since
echinococcosis, trichinosis, diphillobothrosis, opistorchosis and other
helminthoses are found in arctic foxes, the possibility of the trans-
mission of these diseases to humans is not excluded. (A. Ch.).

Genus of [Common] Foxes
Genus Vulpes Oken, 1816

1775 Vulpes. Frisch. Natur-Syst. d. Vierfiiss. Thiere, p. 15. By
decision of the International Committee on Nomenclatural,
this work was considered to have no nomenclatural signifi-
cance.*

1816 Vulpes. Oken.* Lehrb. d. Naturgesch., 3, Th. 2, p. 1033,
1034. Vulpes communis Oken = Canis vulpes Linnaeus.

1839 Cynalopex. H. Smith. Jardine’s Nat. Library. Mamm., 25,
р. 222. Canis corsac Linnaeus (У.Н.).

Species of medium and small size.

Skull light and slender, elongated, with widely situated zygo-
matic arches, and low (height in occipital region less than 1/3 of
condylobasal length). Skull sculpturing is weakly defined, sagittal
crest not developed at all or weakly defined only in posterior part
of the parietal bones. Facial part light and slender, considerably,
sometimes very strongly elongated and much longer than braincase;
its length from the posterior wall of infraorbital foramen to posterior
edge of the canine alveoli greater than skull width above the ca-
nines, while distance from line connecting ends of supraorbital
processes to anterior end of premaxillae significantly greater than
that to upper edge of occipital foramen.

Interorbital (frontal) region flat, not swollen between supraor-
bital processes—here a concavity (groove) is located; supraorbital
processes large, dorsally somewhat concave, with small air cavities,
a quite sharp edge and with apex directed somewhat downwards.
Profile line of rostral part of skull grades very gradually into frontal-
parietal line, not forming a noticeable, or only an ill-defined, projection;
sometimes, upper profile of skull conforms almost to a completely
straight line.

Posterior edge of tooth row reaches level of anterior edge of
interpterygoidal fossa. No deep notch and projection on lower edge

*However, the International Commission on Zoological Nomenclature (1979)
retained Vulpes Frisch 1775, while rejecting Vulpes Oken 1816.

386

of lower jaw in front of angular process.

Dental formula СТР Ма = 42. In some, as individual de-

viation it may be СТР М5 = 40. Teeth relatively small and

light, with very sharp cusps and ridges. Length of upper carnassial
tooth and molars 2%—3 times more than palatal length. Canines thin,
strongly curved and very long—when jaws are closed, tips of upper
canines reach or nearly reach lower edge of mandibular bone or
even extend beyond it, while tips of lower canines are located
above edge of alveoli of upper canines (Fig. 18). Ends of upper
canines directed straight downwards; no cutting edge develops on
posterior side of canine; canine surface entirely smooth. Incisors
usually without accessory lobes.

Trunk long, limbs quite short, tail very long—more than half that
of body, always falls below tarsal joint, and in standing animal
reaches the ground. Head elongated, with long pointed muzzle.
Ears long, when bent forward always reach eye, sometimes very
long with pointed tip. No large bare callosity found on the rear
extremities behind four bare toe pads (a small one occurs—feet
covered with fur). Glandular area is found in front of the middle
callosity on the fore and hind limbs.

In some, fur dense, fluffy, long and soft (northern forms), in
others, more sparse and coarse. Color monophasic, in one species
(red fox, V. vulpes) polyphasic, and, except for black phase of this
species, adults do not exhibit only one color. In various species, the
prevailing colors are reddish, sandy- and clayey-yellow and brown
tones with silvery-white and black in the form of admixtures and
black markings. Supracaudal gland well developed. Nipples 4 or 6
pairs. No marked sexual dimorphism (males are slightly larger), age
dimorphism well manifested and seasonal dimorphism 1$ sharp in
some forms (northern), but is revealed almost solely in degree of
fur density and length. There are two molts per year.

In general appearance, species of the genus are relatively very
monomorphic, differing mainly in size. In this respect, three types
occur: the relatively large red foxes (V. vulpes), forms of lesser
size—corsac fox (V. corsac) and its analogues, and very small
foxes of the Baluchistan type (V. cana). The largest forms have a

268

387

body length up to 90 cm and weight up to 14 kg and the smallest—
about 40 cm in body length and hardly more than 2 kg in weight.

They are inhabitants of extremely varied landscapes—from tun-
dras and taiga to extremely high mountains, and steppes and deserts.
They prefer areas with more or less open expanses and avoid the
dense thick forests. There are no regular migrations. They are
monogamous, uniting for one reproductive season. Up to 13 young,
usually less, born in burrows. They are carnivorous animals, feeding
on small vertebrates, mainly rodents, carrion, and as an exception,
on insect and plant food. Except for the breeding season, they lead
a solitary way of life.

The range of the genus is very vast, located on four continents
and in all climatic zones. It occupies all Europe, to North Cape and
islands of the Mediterranean Sea, includes the British islands (ab-
sent in Iceland), all Africa except Madagascar, and Asia northward
to the northern shores of the continent (except the northern Taimyr)
and the Kolguya islands, Yuzhnye Novaya Zemlya and Belyi is-
lands (locally wandering animals may be seen farther north; see
description of red fox, V. vulpes) and southward to Tonkin, Yunnan,
Sichuan, Assam, Bengal and India to Cape Comorin. It occupies
the entire Arabian Peninsula. The range includes Karagin, St.
Lawrence, Kuril, Sakhalin, Shantar and Japanese islands. Species
of this genus are absent on the islands of the Bering Sea (except
those mentioned).

In North America, the range extends northward to the northern
coast of Labrador, Hudson Bay, a line from Chesterton (on the
western shore of Hudson Bay) to Coppermine (on the Arctic coast),
and farther westward on the Arctic Ocean and Bering Sea coasts.
Southward, the range extends to California and northern Mexico
inclusive, but does not include the southern and southeastern states
of the USA from Maine to Louisiana (for details, see range de-
scription of red fox, V. vulpes). The range includes Newfoundland
and Kodiak islands. Species of the genus are absent from the
coastal belt of British Columbia and adjacent islands. Acclimatized
in Australia (V. vulpes). In the vertical direction, they reach ex-
treme heights. The range of the genus was not subjected to
substantial reduction in the past century.

In systematic characteristics, Vulpes represents a more spe-
cialized group than Canis and Alopex, and the degree of specialization
of some species (V. ferrilata, V. cana) is. very high. Besides the

389

close relationship to the arctic fox, Alopex (see above), Vulpes is
closely related to the fennec group—genus Fennecus. All three
genera are considered “neighboring” groups. Through Alopex,
Vulpes is related to Canis (wolves and jackals) (see characteristics
of family and of named genera). The relation to genus Ducicyon
(American gray fox and close species), with which the described
genus is sometimes associated is, apparently, more distant. Since
the extreme forms of all the genera mentioned have much in com-
mon, and serve as connecting links between them, there is basis for
considering all of them as only subgenera of one broadened genus—
Canis (see the family characteristics). At the present time, however,
it is still almost always considered as qualifying as independent.

The genus already was separate in the Pliocene and may have
appeared even in the lower Miocene (North America). In Europe
and Asia, it is known in the Pleistocene. The genus Vulpes, like
Canis, probably developed from the American Mio-Pliocene group
Tomarctos.

In the contents accepted here, the genus 15 quite monotypic and
was not subjected to any considerable subdivision. Attempts were
made to split into a separate genus the corsac fox (V. corsac) and
unite it (sometimes also with V. velox) with genus Alopex (arctic
foxes).

In the genus are 6 species: 1) red fox, Г. vulpes; 2) corsac fox,
V. corsac; 3) Bengal fox, V. bengalensis; 4) Afghan fox, V. cana;
5) American small fox, or American corsac, V. velox* and 6)
Tibetan fox or Tibetan corsac, V. ferrilata. The genus includes
abou. 20% of the species of the family, 4 Asiatic species (V.
corsac, Г. bengalensis, Г. cana, Г. ferrilata), 1 American spe-
cies (Г. velox)* and 1 species distributed in the greater part of
North America, in almost all Eurasia and in North Africa (V. vulpes).

The African Г. chama and the African-Near Eastern Г.
rueppelli are sometimes included in genus Vulpes; however, they
are apparently to be referred to genus Fennecus.

The fauna of the USSR includes 3 species: 1) red fox,
V. vulpes Linnaeus, 1758; 2) corsac fox, V. corsac Linnaeus,
1768; 3) Afghan fox, Г. cana Blanford, 1877, i.e. 50% of the

*In English, swift fox; V. megalotis, the kit fox, is often considered a distinct
species as well—Sci. Ed.

269

390

species of the genus, and about 1% of the species of the USSR
fauna'.

The range of the genus covers the entire territory of the USSR,
except the above-mentioned northern regions. Species of the genus
are met with in all landscapes of the country.

The members of this species are useful as commercial fur-
bearing animals, and in part for limiting the number of harmful
rodents. In places, they cause some harm by killing useful sporting
animals and birds (V.H.).

AFGHAN FOX?
Vulpes cana Blanford, 1877

1877. Vulpes cana.* Blanford. Journ. As. Soc. Bengal., 46, pt. 2,
p. 231. Gwadar, Baluchistan (Mekran coast).

1907. Vulpes cana var. nigricans. Shitkow. Zool. Anz., 32, p.
448. Southern Tadzhikistan (“Gornaya [montane] Bukhara’).
V.H.

Diagnosis

Dimensions very small, body length not more than 50 cm,
condylobasal length of skull not more than 91 mm. Tail tip not
white, back color gray mixed with black ripples, without reddish and
brown tones, or they are very weakly developed (V.H.).

Description

The Afghan fox is a carnivore of typical “foxy” appearance, but

of very small size. It has relatively short legs, elongated trunk and

very long tail. Its length is considerably more than half the length

of the body, and in a standing animal it reaches the ground. Head

with very pointed, typically fox-like muzzle and large, wide, erect

pointed ears. Their length more than half the length of hind foot.
Number of teats unknown.

"Information on the occurrence of V. rueppelli Schinz, 1825 [“И. (Megalotis)
famelicus Cr.”; Flerov, 1932] in southern Turkmeniya is a mistake.

2Shardel’” of furriers.

* Misspelled “canus” in Russian original—Sci. Ed.

270

Fig. 101. Afghan fox, Vulpes cana Blanf. Sketch by A.N. Komarov.

In winter, the fur is dense, fluffy, very soft and relatively quite
long. Dorsal guard hairs 45—65 mm long, underfur, 35-45 mm. Tail
covered with dense, long hairs (their length 65—90 mm) and very
fluffy.

General tone of winter fur brownish gray, with well-developed
black highlights along back originating from guard hair color. Guard
hairs have black terminal portion, with a small silvery-white zone
proximal it. Some guard hairs are entirely black. Guard hairs form
on the skin a silvery frosting. On head, general color lighter and
somewhat whitish. Well-defined blackish-brown stripe runs from
corner of eyes to base of vibrissae. Light rusty frosting develops on
occiput. Rear sides of ears gray in color or blackish-gray. Chin
dark brown or brownish-black. On sides of the body, color consid-
erably lighter than on dorsum, changing into pale yellow-gray. Here
admixture of black guard hair is small; guard hairs have gray and
silvery tips and underfur lighter than on back. Throat, lower part of
neck, chest, and whole abdominal surface white, with light straw-
colored tones. Extremities gray in color, more or less dark, with
light rusty highlights on external side. On fore feet, black markings

392

sometimes present. Tail gray, with strong blackish highlights above.
In region of supracaudal gland, on upper side near base, is a well-
defined dark spot. Underfur is mouse-gray in color, their tips with
light, or more or less intense ocherous highlights.

Individual variation in the color described above are known. It
is expressed chiefly in that individual specimens are tinged espe-
cially dark gray in color without any ocherous or rusty admixture
or with a very weak admixture of these tones, above all on the legs.
In others, the admixture of ocherous and rusty (“foxy”) tones is
more noticeable, and the rusty is well expressed on the shoulders.

Summer pelage is not described. The first [juvenile?] pelage
and age variation in color are not known.

The fox we are describing is very rare in collections and its
amplitude of variation is poorly known. Generally, this species is, in
many respects, poorly studied.

Skull resembles that of corsac fox, but facial portion relatively
shorter, braincase relatively larger and more expanded. Zygomatic
arches widely separated and are quite broad anteriorly, from which
its general outline is more or less rounded (together with rostral
portion, not forming evenly tapered clinoidal figure). Nasal bones
narrow and highly tapered towards posterior end. Sagittal crest not
developed, and temporal lines stand far apart from each other,
forming a lyriform figure. Molars relatively large; canines thin and
long. Middle incisors without accessory blades.

Fig. 102. Skull of Afghan fox, Vulpes cana Blanf. (after Pocock, 1941).

271

393

Chromosome number unknown.

Size very small—this is the smallest species of the genus in our
fauna, and in the genus generally. Body length 40—50 cm, tail length
33-41 cm, hind foot about 100 mm, ear length about 90 mm.

Condylobasal length of skull, 89-91 mm; zygomatic width, 50—
51 mm; interorbital width, 18—19 mm; length of hard palate, 45 mm;
length of nasal bones about 29 mm, length of upper molar row, 41—
44 mm; length of lower jaw, 67-69 mm; length of upper canine
about 10 mm; and of lower, 8 mm (according to Pocock, 1941, and
Novikov, 1956; published data are those of separate individuals; in
our museums, there are no fully valuable materials).

Information on comparative sexual dimorphism in size is absent;
as in all foxes, females are probably smaller than males (V.H.).

Geographic Distribution
Eastern part of Near East.
Geographic Range in the Soviet Union

This represents the extreme northern periphery of the species range.
In dimensions it is insignificant and is limited to the extreme south
of Middle Asia.

Reliable places of capture of the Afghan fox lie in the Kopet-
Dag at the northern foot of the western part of the Sarymsakli
range, 40 km from Germab, which is west of Ashkhabad (1926;
Ognev, 1931), and at Bakharden on the railway line between
Ashkhabad and Kyzyl-Arvat (1926; Novikov, 1956).

In the 90’s*, 4 skins were brought to Moscow from
“Vostochnaya [East] Bukhara’, 1.е., southern Tadzhikistan (Zhitkov,
1907); one of these specimens is preserved in the Zoological Mu-
seum of Moscow University in stuffed form. At that time, the
possibility was admitted that these were not native, but imported
from Afghanistan. The majority of later authors shared this belief.

At the present time, information has appeared that skins of this
species occasionally come in from southern Turkmeniya and
Tadzhikistan (Kuznetsov, 1952). According to other data
(Chernyshev, 1958), “during the course of a prolonged period”, not

*In Russian original, “У 900-kh godakh” [In the 900’s; i.e. 10th century].
This evidently refers to the 1890’s—Sci. Ed.

272

394

once has a skin of this species appeared among the fur prepared
in Tadzhikistan. Thus, the question on the occurrence of Afghan
foxes on the eastern shore of the Amu-Dar’ya and Pyandzh re-
mains open and, most likely, will be settled in the negative.
Southwestern Turkmeniya remains the established area of its oc-
currence within the borders of our country.

Apparently, one must now reach the conclusion that the
Afghan fox in our country is only vagrant, appearing rarely in the
Kopet-Dag and adjacent plains. This corresponds to the known
occurrences of the species in northeastern Iran (Khorassan; Duruh
in the eastern part of the province). It may be assumed that the
species under description appears also in southeastern Turkmeniya—
along the Tedzhen and between Tedzhen, Murgab and Amu-Dar’ ya
[rivers]. These areas lie south of the places of transience of the
Afghan fox to the west [Fig. 103]. Wandering into Tadzhikistan is
also doubtful because the Pyandzh and Amu-Dar’ya are probably
insuperable barriers for such a small animal.

Geographic Range outside the Soviet Union

This includes Afghanistan, the northwestern provinces of India
bordering Afghanistan, Sind, Baluchistan and eastern Iran, in par-
ticular Khorassan.

The range of the Afghan fox is notable for its very small size,
which is unusual for a carnivore, and particularly for the species of
the family Canidae. It is even smaller than the range of the rela-
tively narrowly distributed Bengal (V. bengalensis) and Tibetan (V.
ferrilata) foxes (V.H.).*

Geographic Variation

Geographic variation in this species is not described, and probably,
is not manifested—its range is too small for this. Moreover, it is
restricted to one natural geographic landscape region. Based on all
data, the above-mentioned specimen described from “Vostochnaya
Bukhara” as “var. nigricans” represents only an individual color
deviation (V.H.).

*V. cana has recently been found in northwestern Iran, the Arabian peninsula
and Israel (Harrison and Bates, 1991), where it occupies steep, rocky habitats—
Sci. Ed.

271

273

395

Fig. 103. Range of the Afghan fox, Vulpes cana Blanf., in the USSR. Dots
designate places where animals were obtained, question marks—areas
of assumed occurrence. V.G. Heptner.

Biology

There is no information whatsoever about the biology of the
Afghan fox within the USSR. Data from other countries where this
species lives, especially India and Pakistan, are also absent (V.H.).

Practical Significance

In our country, the Afghan fox has no practical significance at all.
As for the small number of skins, “зВагае! ”, which sometimes
appear among lots of furs from Tadzhikistan and Turkmeniya, and
probably from the neighboring countries (see above) they do not
play any role whatsoever in fur trade. Apparently, the agricultural
significance of this species is also negligible. No substantive mate-
rials are available from the other countries (V.H.).

396

272 . Fig. 104. Species range of the Afghan fox, Vulpes cana Blanf. Very schematic.
V.G. Heptner.

CORSAC FOX

Vulpes corsac Linnaeus, 1768

1768. Vulpes corsac. Linnaeus. Syst. Nat., Ed. XII, 3, App. p.
223. Steppes between Ural and Irtysh rivers. Restricted to
“Pri-Irtysh steppes south of Petropavlovsk” (Ognev, 1935).

397

1912. ? Vulpes corsac var. nigra. Kastschenko. Kashchenko.
Ezhegodnik Zool. muzeu I. [Irkutsk] Akad. Nauk, 17, p. 393
Trans-Baikaliya.

1935. (1934) Vulpes corsac scorodumovi. Dorogostaiski.
Dorogostaiskii. Izv. Irkutsk. gos. protivochumn. inst. Sibirii i,
D.V. kraya. 1, р. 47. Southern Trans-Baikaliya.

1935. Vulpes corsac kalmykorum. Ognev. Zveri SSSR 1 prilezh.
stran [Animals of the USSR and adjacent countries]. 3, p.
634. Kalmytsk steppes (steppes on right bank of Lower
Volga; V.H.).

1935. Vulpes corsac turcmenicus. Ognev. Zveri SSSR i prilezh.
stran [see above] 3, p. 635 “Turkmen and Tadzhikistan
semideserts and deserts’. It is proposed to restrict this to:
Murgab and Tedzhen districts and lowlands at foot of east-
ern Kopet-Dag (east of Ashkhabad). V.H.

Diagnosis

Of moderate size, larger than Afghan, but smaller than red foxes.
Body length more than 50 cm but less than 65 cm, basal length of
skull more than 95 mm, but less than 112 mm. Color grayish-
ocherous with gray hairs, back side of ears not black, end of tail
not white (V.H.).

Description

In general construction, the corsac is a typical fox and very similar
to the red fox, representing itself as a reduced copy. It is also
stocky and has an elongated trunk,* but the tail is not nearly so
fluffy and is relatively somewhat shorter. Its length constitutes half
the length of the trunk or slightly more; in the standing animal, it
reaches the ground. The corsac has relatively somewhat larger
ears; they have broad bases and pointed tips.

Number of teats unknown.

Winter fur dense, soft and silky. Its general color is pale-yellow
gray with ocherous tints. Along the middle of the back, a noticeable

On title page, designated 1934, on cover, 1935. Appeared in 1935 (Signed
by press).

4The idea that the corsac has longer limbs than the red fox (Ognev, 1931) 1$
untrue.

274

398

brownish tone and well-developed “silvery”, resulting from silvery-
white ends of guard hairs. Strongly developed silvery tint sometimes
makes fur of the animal silvery-gray on the back. Sometimes, on
the contrary, brownish tone quite strongly developed. Shoulders
same colored as on dorsum. Side also with same tone, but consid-
erably lighter. Lower surface of body, including chest and inguinal
region, white or with light yellowish frosting. Fore legs light-yellow
anteriorly, laterally rusty-yellowish; posteriorly, same color but paler.

Head grayish-ocherous or brown in color, darker on forehead.
Around eyes color lighter, a dark field of triangular form between
anterior corners of eyes and upper lip. Around mouth, on throat and
along lower neck, color white with weak yellowish tint. Rear side
of ear ocherous-gray or reddish-brown, bordered anteriorly along
edge of ear in white, interior of ear pinna covered with yellowish-
white hairs.

Tail covered with dense, fluffy hairs, and has a dirty dark-
ocherous or gray-brownish general tone. On dorsal side it has blackish
tinge; on basal half—6—7 cm from base in region of supracaudal
gland, is found a dark spot. At base, brownish tone noticeable.
Lower side pale. Tip of tail dark, usually almost black. Underfur

Fig. 105. Corsac fox, Vulpes corsac L. Sketch by A.N. Komarov.

PT Ци:

$ 1:
В =
Е

bah AT
7

—

› 277

ея -
к W IN te
ie < \\ NIN

‘ S
>
ee |

55 eg. << 6
ее
СХ

NS : |
Meant AN | `
м. WwW woe Hh,

к у
AWN Sar

«<

275 Fig. 106. Skull of corsac fox, Vulpes corsac L.

399

277

400

ash-gray, brownish at ends or slightly rusty. Guard hairs gray at
base, white at tip, pale-brown in middle. Individual guard hairs have
two dark zones with an intermediate white one.

Summer fur differs greatly from winter—tt is short, sparse and
coarse; tail hairs are also considerably shorter. In this fur, the
corsac fox looks wiry and long-legged, with a large head. General
color is more even—back and sides are nearly undifferentiated,
dirty-ocherous or dirty-sandy, dull. Gray hairs are not developed in
summer.

There is no sexual dimorphism in color.

In general, the skull of the corsac fox resembles that of the red
fox, but, irrespective of the smaller dimensions, differs from it in
many essential characteristics. In general appearance the skull is
much shorter and broader and not at all elongated. The general
outlines of the zygomatic arches and rostral region are reminiscent
of a broad wedge—zygomatic arches are very broad posteriorly,
and the skull in the region of the infraorbital foramen is broad, and
not as compressed as in the red fox. Therefore, the line of the
zygomatic arches merges into the line of the rostrum more gradu-
ally, without such a break as in the red fox. The rostral region is
relatively short—the ratio between the length of the nasal bones and
the length of the distance between their posterior end and the
posterior-most point of the occipital region, is 1: 1.6—1.8.

The braincase protrudes behind the transverse line passing
through the articular condyles of the lower jaw for a shorter dis-
tance than in the red fox. The general volume of the quite convex
and swollen braincase is relatively greater than in the red fox. It is
comparatively short, but widened posteriorly. The line of the upper
profile is straighter. The occipital crest is weakly developed, the
sagittal, very poorly and 10—15 cm in length. The temporal lines
come off of the postorbital processes almost parallel, and very

‚ gradually narrow posteriorly, uniting with the sagittal crest. They

outline a wide lyriform or triangular expanse on the frontal and
parietal bones. Orbits are relatively large—their greatest height is
equal to or more than the width of the interorbital region. Paraoccipital
processes are weaker than in the red fox—their ends do not reach,
or only just reach, the level of the lower edge of the auditory
meatus. The length of the lower jaw symphysis is 4.5—6.5 times
shorter than its greatest length.

40
80

a
160 170
of movements. Dot refers to

we
A

к
ААУ

nd place

ad). V.G. Heptner.

т
SI] -

|=
=
|
|=
|

L. in the USSR. Arrows show direction a

fox, Vulpes corsac
of especially long movement in the west (Pavlogr

place

276 Fig. 107. Distributional limits of the corsac

402

Canines and carnassial teeth are comparatively stronger than in
red fox. When the jaws are closed, the tip of the upper canine
reaches the lower edge of the mandibular bone, the apex of the
lower canine extends beyond the edge of the alveolus of the upper.
Sometimes, as a form individual variation (age?), reduction of the
second lower molar is observed and the dental formula becomes

Е РМ =n.
By A pee)

Other than size, there are no sexual differences in the skull.

Number of chromosomes unknown.

Only approximate body size of the corsac fox is known, since
the data published are those of individual measurements of animals
“in the flesh’. Data on the skull dimensions are somewhat more,
but the material is very uneven.

Body length, 50—60 cm; tail length, 25—35 cm; height at shoulder,
about 30 cm; length of ears, about 8 cm; length of sole of hind foot,
about 11 cm.

Greatest length of skull, 102.2—124.0 mm; condylobasal length,
98—118.1 mm; zygomatic width, 57.5—71.1 mm; skull height in region
of auditory bullae, 39.0-45.7 mm.

Sexual dimorphism in size is almost not expressed, and appar-
ently males average only slightly larger than females. The belief
sometimes reported, that females are larger than males, is incorrect
(V.H.).

Geographic Distribution

Flat steppes and semideserts, in part deserts of Asia and Southeast-
ern Europe.

Geographic Range in the Soviet Union

This is relatively small, but constitutes a considerable part of the
western range of the species. It occupies the steppes and deserts
of the southeastern European part of the Soviet Union, northern
Caucasus, Kazakhstan, Middle Asia and Trans-Baikaltya. It is di-
vided into two parts isolated from each other, uniting outside the
borders of the USSR.

The main and vastest part of the range occupies Middle Asia,
Kazakhstan and adjacent territories. In the west, the southern

278

277

278

403

borders of the distributional area of согзас fox includes the Cis-
Caucasian steppes on the south. Beginning on the Caspian coast at
Makhachkala, it passes along the foothills to Khasav’yurt, farther
to Groznyi and Mozdok, and then, approximately to Stavropol’ and
from there, embraces the Egorlyk basin, and passes on to the lower
Western Manych and to the lower Don [rivers].

Along the Don, which constitutes the western border, it rises
approximately to 39° М. Lat. (somewhat north of Volgograd)°. Thence,
the line of the border, following the left bank of the Volga, reaches
northward almost to the Samarsk bend, and then ascending somewhat
more to the north and to the east and becoming the northern border
of the range, it encloses the extreme southern regions of the eastern
half of the Tatar Republic (Bugul’minsk, Bavlinsk, approximately at
54° 30'N. Lat.) and adjoining—Bashkirtya. Farther, it passes along the
outliers of the Bashkirian steppes and along the Obshchii Syrt it turns
sharply to the south to the southern end of the Urals.

Rounding the southern spurs of the Ural mountains along the
right bank of the Ural [river], the border on the eastern slope of the
range goes northward and passes through Verkhnaural’sk. Thence,
going eastward through Troitsk, Shadrinsk, Tyumen’, Taru, Ishim,
Vengerovo, Barabinsk, Omsk and Lake Chany, the boundary line
passes through the bend of the river at Kamen’ on Ob’. Following
the left bank of the [Ob’] river, it descends to Barnaul, rounds the
Altai from the west and along the northern slope of the Zaisan
depression, exits into China.

In Middle Asia and Kazakhstan, the range occupies all the level
expanses, i.e. corsac fox is absent in Tarbagatai, Dzhungarskii Alatau,
in the entire Tien Shan (occurs in the Karatau) and Pamir-Alai and

‘There is not sufficient exact information about the normal occurrence of
corsac fox to the west of the Don. Therefore there is no basis at present to assign
the western border of the range along the “eastern part of the Dnepr-Don
interfluve (Pidoplichko, 1951). At the same time, it is probable that some centuries
ago, when no cultivation was present in the southern European part of the Soviet
Union and people were very scarce, the corsac fox lived somewhat farther west
than now; probably it lived normally in the Priazov steppes. However, positive
information on this is absent.

It is not excluded that there were intrusions farther to the west as far as
Zaporozhsk district and western Priazov. However, intrusions to Zaporozhsk
district were extremely rare (Sharleman’ [Charlemagne], 1937). There is no basis
for believing that corsac fox normally lives and is even sufficiently common (at
Korsak-Mogila; Sharleman’ [Charlemagne], 1948).

279

404

Kopet-Dag. It is found in the level parts of the right bank of the
upper Amu-Dar’ya in Tadzhikistan, but it is, apparently, absent in
the Fergana valley. From Turkmeniya, where the mountain range
(Kopet-Dag) forms no obstacle, the range passes to Afghanistan
and Iran, and from eastern Kazakhstan, through the Dzhungarsk
Vorota [Gate] and Zaisan depression, into Dzhungariya.

Another section inhabited by corsac fox is found in our country
in southern Trans-Baikaliya. This region represents the northern
outlier of the Manchurian and Mongolian part of the range. In the
adjacent USSR, the range is separated from the already-described
western part. The Trans-Baikal section of the range is very small
and, in its turn, is divided into two parts. The first includes only the
steppes along the Argun’ and Onon, and the Aginsk steppe (the
area lying to the west of the lower course of the Onon). The
second section is still smaller and occupies the steppes along the
Selenga and Dzhida in western Trans-Baikaliya.

The northern border of the main range of corsac fox cannot be
considered sufficiently defined. Beside the insufficient study of the
corresponding territories, the reason for this is that corsac fox is
characterized by more or less regular seasonal, and also irregular,
migrations. Moreover, through the force of confluence of various
ecological conditions, considerable fluctuations exist at places in its
numbers, especially at the borders of the range. Finally, the corsac
fox under harvest very easily yields to extermination, and slowly
restores its numbers. Therefore, in individual years or periods it
may, firstly, shift somewhat northward the northern border of its
reproductive area or, secondly, there may occur locally quite long
transgressions by stray individuals.

At the same time, due primarily to limited information about this
species, it is essentially impossible to firmly delimit breeding terri-
tory, and territory of migration. The outlined northern border is, to
а. considerable degree, determined by more or less regular, if none-
theless rare, migrations, and the true limit of the breeding area lies,
apparently, somewhat southward. Thus, in southern Tatar Republic,
the corsac fox occurs only as a transient (migrating), and the actual
[northern] limit between the Volga and Ural apparently lies at the
latitude of the Samarsk bend (about 53°). The same thing applies
to the extreme northern points where corsac fox is encountered
between the Don and Volga. In exactly the same way, the outlined
southern and southwestern border in the Cis-Caucasus is, to a

280

405

considerable extent determined by migrations. The area of normal
occurrence here is restricted (or, until recently was restricted) to
the semideserts to the north of the Terek; in the west it did not
reach Stavropol’.

Long migrations are known for the corsac fox from the steppes
in the southern Ural foothills to Zilair and in Shadrinsk region (at
56°), from the Privolga steppes westward to Hel’govk (about 50
km to the north of Nogaisk on the shore of Berdyansk gulf), and
to Pavlograd (east of Dnepropetrovsk). Within the range, there are
sections where the corsac fox is absent (Ulutau, Kazakh upland) or
they occur as transients (southern Pribalkhash) (see section on
“Biology”).

Geographic Range outside the Soviet Union

This occupies the steppe part of northeastern China, in all events
Barga (expanse between the Argun’and Great Khingans), and prob-
ably the Manchurian plains in the Nonn’ and Sungari basins; Inner
Mongolia; the entire Mongolian Republic, except the taiga districts
of the Kentei mountains, the wooded and mountainous districts of
the Khangai [mountains], and the Prikosogol’sk district*; Dzhungariya
and Kashgariya; Afghanistan (probably only northern regions); and,
apparently, northeastern Iran.

The limits of the distribution of the corsac fox in southern
Central Asia is not known—it is possible that it extends to the
ranges that delimit Tibetan Plateau on the north (see section on
“Geographic Variation”).

Therefore, as a whole, the range consists of two vast areas,
western and eastern (central-Asiatic), with a narrow “isthmus”
uniting them in the area of the Dzhungarsk Gate and Zaisan de-
pression.

In the Pleistocene, corsac fox was widely distributed in Middle
and, apparently, Western Europe and it still lived in the Crimea in
the Holocene® (V.H.).

*Area around Hobsogol Lake—Sci. Ed.

®Geographic distribution of the corsac fox according to Dinnik, 1914;
Ognev, 1931; Shergin and Frenkel’, 1936; Sharleman’, 1937, 1948; Heptner and
Formozov, 1941; Heptner et al., 1950, V.A. Popov and Lukin, 1950; Shvarts,
Pavlinin and Danilov, 1951; Pidoplichko, 1951; Kirikov, 1952; Yanushevich and
Blagoveshchenskii, 1952; Afanas’ev, Bazhanov et al., 1953; Bannikov, 1954;
Vereshchagin, 1958; Sludskii, 1959 and a series of other sources and on unpub-
lished material of V.G. Heptner.

279 Fig. 108. Species range of corsac fox, Vulpes corsac L. V.G. Heptner.

Geographic Variation

The geographic variation is quite evidently manifested, although,
apparently less than is sometimes considered (Ognev, 1935).
Within the limits of the USSR, one may differentiate the follow-
ing subspecies.
1. Kazakhstan corsac fox, Г. corsac Linnaeus, 1768 (syn-
onym kalmykorum).

281

407

Relatively large form, with fluffy and dense winter fur of above
described color, without admixture of reddish tones in winter fur.

Greatest length of skull 103—111 mm, basal length 96-105 mm,
zygomatic width 59-68 mm, greatest skull width 37—42 mm, skull
height in the region of tympanic bullae 42.644 mm, length of upper
tooth row 48—55 mm (less than 10 individuals; Ognev, 1931).

Northern part of range of the species, to Cis-Altai steppes. Distri-
butional limits in the south are not established—apparently, it does not
extend farther than the latitude of the northern end of the Aral Sea.

Not outside the USSR.

2. Turkmenian corsac fox, V. c. turcmenica Ognev, 1935.

Considerably smaller than the preceding (smallest form of the
species), with shorter, coarser winter fur, of dirty-grayish color and
reddish tones.

There are no data on body measurements.

Skull measurements of Turkmenian corsac fox (14 males and
females; V.G. Heptner) are as follows (Table 30).

Inhabits the southern part of the species range—plains of Middle
Asia and Kazakhstan, probably, south of the parallel on the Aral
Sea’s northern ‘end. The boundary with respect to the preceding
form is not elucidated—apparently, there is a wide zone of transition
between each other.

Outside our borders—in northern Afghanistan and northeastern
Iran.

The-corsac fox of the Volga-Ural steppes and right bank of the
Volga apparently has some characteristics of transitional character
between the normal form and turcmenica. They are described as
kalmykorum, and their furs are considered as a particular Astra-
khan type.

Table 30. Skull measurements of Turkmenian corsac fox (in mm)

Data Minimum Maximum Average
Total length of skull 102.2 114.0 105.8
Condylobasal length 98.4 110.3 103.0
Length of nasal bone along midline Sei 36.3 33.9
Zygomatic width 57.5 65.3 60.4
Greatest width of cranium 40.7 44.1 42.5
Height in region of tympanic bullae 39.3 43.2 41.6
Length of molar tooth row 38.3 42.5 40.2

Interorbital width 18.4 23.0 20.7

408

3. Trans-Baikal согзас fox, V. с. scorodumovi Dorogostaiskil,
1935 (synonyn nigra).

There are no data on body measurements.

Skull dimensions of animals from southern Trans-Baikaliya, are
as follows (Dorogostaiskii, 1935) (Table 31).

Table 31. Skull measurements of Trans-Baikal corsac fox (in mm)

Data Males (41 spec.) Females (37 spec.)

Minimum Maximum Average Minimum Maximum Average
Greatest length 110.9 124.0 1170. 1102 119.6 114.6
Condylobasal length 106.0 118.2 1135 1060 ТЯ. 0110.5
Zygomatic width 63.0 Ты 66.9 63.0 68.8 65.2
Width above canines 18.4 25 19:9 18.0 21.5 19.6
Maximum width of 42.0 46.1 44.2 41.4 47.7 43.5

cranium

Length of nasal bone 34.0 43.0 38.0 35.0 40.4 37.0
Height in region of 41.2 45.7 43.7 41.5 45.0 43.1
tympanic bullae

Found in Trans-Baikaliya.

Outside the USSR, in the Mongolian People’s Republic and
northeastern China.

To this form, by priority the name nigra Kastschenko, 1912
should be assigned; it is based on a skin (without skull) without
accurate locality reference, but which apparently originates from
Trans-Baikaliya or the contiguous part of the Mongolian Republic.
However, this name is hardly acceptable, since the author de-
scribes an ordinary melanic, and, due to the defective skin, he is not
even sure that it is assignable to this species.

* * *

Separate forms of the corsac fox have not been described from
that part of the range lying outside our territory. One may suppose
that there, except perhaps in Kashgariya, the same subspecies live
which are found in our country.

In the western literature, there is a widely distributed idea (see,
e.g., Ellerman and Morrison-Scott, 1951) that V. eckloni, described
by Przheval’skii (1883) from northern Tibet, constitutes a subspe-
cies of V. corsac. This idea is mistaken—V. eckloni is a synonym
of V. ferrilata. The error is probably based on the fact that in

282

409

earlier works (in part, as indicated), Przheval’skii called this species
the corsac fox (V.H.).

Biology

Population. In the steppes of the Pri-Black and Pri-Azov Seas,
corsac fox became extinct, apparently, only in the 18th—19th cen-
turies. The cause for this was believed to be the ploughing of the
virgin steppes (Pidoplichko, 1951; Topachevskii, 1956), but, appar-
ently, its extinction took place under the influence of other
factors—climatic deterioration (it became less continental) and
others. Reduction in numbers of the corsac fox also may have been
influenced by the extermination of the steppe marmot [Marmota
bobac], the burrows of which it occupies particularly eagerly, and
by massive reproduction of the red fox in cultivated fields, repre-
senting not only its competitors but also its enemies.

At the beginning of the current century, corsac fox was not
rare in the Don steppes, in Ergeni [hills] and in the Astrakhan
semideserts and south of the latitude of Volgograd (Artsibashev,
1859; Dinnik, 1914; Kirikov, 1959). In recent years, its population
even increased, and in Ergeni the animal is now common.

In the Cis-Caucasus, this carnivore is common in the region of
the Kumo-Manychsk canal and in the Kum-Terek interfluve. Dur-
ing this century, in the latter region, its population was subjected to
significant changes. Thus, in the beginning of this century, in the
northern Caucasus, it was communicated (Dinnik, 1914) that: “Pre-
viously, in all these very poorly settled places, corsac foxes lived in
great numbers, but with settlement of the steppes and their plough-
ing, these animals began to greatly decrease in numbers; in many
places they even became completely extinct’. In 1924—1925, corsac
foxes again became numerous in the northeastern Cis-Caucasus. A
single hunter caught 15—30 animals in one season (Ognev, 1931;
Heptner and Formozov, 1941). By 1931, the population of this car-
nivore was again strongly reduced. There were relatively many
corsac foxes in the Cis-Caucasus in 1951 (Vereshchagin, 1959).

East of the Volga on the Obshchii Syrt and adjacent regions
northward to the upper Bol’shoi and Malyi Irgiz rivers, the corsac
fox periodically appears common, since its skins are obtained by the
hundreds (N. Severtsov, 1861; Zarudnyi, 1897; Bazhanov, 1930;
A.A. Sludskii). It is very rarely found in the steppes of Bashkiriya,

283

410

penetrating in occasional years to the Beloretsk region and southern
Tatariya (Popov and Lukin, 1949; Shvarts et al., 1951). At the
southern end of the Ural mountains, the corsac fox becomes more
common, for example, in Orenburg district, along the border with
Kazakhstan. In the Volga-Ural interfluve, the corsac fox is again
frequent and, for example, in the Ural’sk district in different years
up to 5,500 individuals are caught in a season, while in Gur’evsk
district—up to 1,700. To the south, on the Mangyshlak Peninsula
and Ustyurt Plateau, it is common (up to 500 spec. caught per
season). It is also common in Mugodzhar.

In western Siberia, the corsac fox appears in various years in
the northern forest-steppe zone and even on the edge of taiga
forests, but here, its numbers are extremely low. In the southern
half of the forest-steppe zone (Barabinsk and Kulundinsk steppes),
the corsac fox is rare and not a permanent occupant. In the south-
ern part of the steppe zone of western Siberia (Chelyabinsk,
Kustanaisk, Severo-Kazakhstansk, and Pavlodarsk districts and Altai
territory), the carnivore being described was common in the last
two decades and in some places, it was abundant. In separate
years, in each of these districts, from 1000 to 3000 skins were
obtained.

The abundance of corsac foxes in typical steppes may be judged
by the number caught by hunters. In years with high populations of
corsac fox, in central Kazakhstan, hunters catch 20—50 and more
animals in a season. In 1947, in Tselinogradsk district, 37 burrows
of corsac fox were found in an area of 30 km’, and of these 8 had
litters, in 8 burrows there had been, apparently, litters, and 9 were
inhabited by single animals. In 1948, in this same region, 34 burrows
were found, of which only 4 had litters (Chirkova, 1952). It is also
numerous in the semideserts of central Kazakhstan (Tselinogradsk
and Karagandinsk districts), and also in the Pri-Irtysh steppe
(Semipalatinsk district). In some years, in central Kazhakhstan, up

to 4,500 corsac skins were obtained in a season. In the semidesert,

the corsac fox periodically becomes very abundant. Thus, near the
Kulanutmes river from July 29 to August 5, 1951, 27 burrows of
this animal were found in an area of 1,000 hectares, of which 7 had
litters in spring (A.A. Sludskii). In years of massive reproduction,
some hunters capture up to 100—120 animals in one season. The
corsac fox is abundant from time to time in the Pri-Aral’sk Karakum,
common, especially in winter, in Betpak-Dala, occurs in the Pri-
Chuisk Muyunkum and in northern Pri-Balkhash’.

411

In the sandy deserts south and east of Betpak-Dala-Muyunkum,
Taukum and Sary-Ishikotrau the corsac fox, as a rule, does not live
permanently, but comes from the north only in certain severe win-
ters. This animal is extremely rare in the north and west of the
sandy desert of Kyzylkum (Bogdanov, 1882). In the Amu-Dar’ya
delta, it is rare (Salikhbaev, 1950). In Kara-Kalpakiya, from 1956
to 1959, only 21 to 76 skins were prepared annually. In the Syr-
Dar’ya valley, on the clayey plain Dar’ yalyktakyr lying to its north,
and in the sands of the Pri-Aral’sk Karakum, the corsac fox is not
especially rare, as in the clayey sections of the Ustyurt. There also,
on sands, for example Asmantai, they are scarce. In the Ustyurt
and in the northern Pri-Caspian, corsac foxes in autumn and winter
time are concentrated on the seashore and near the abandoned
summer camps and villages, where they feed on refuse and
rodents.

In Turkmentya, it is “very common” in many regions (Radde
and Walter, 1889). Thus, it is common on the southern border of
the Karakum and in the Badkhyz (Heptner, 1956). At the same
time, in the Krasnovodsk region, this carnivore is scarce
(Vinogradov, 1952). In the 1924/25 season in Turkmeniya, 11,522
skins were tanned and in 1925/26—7,174 skins (Kogan, 1931). From
1948 to 1959, the number of corsac foxes caught here ranged from
694 (1948) to 2,298 (1956) per year. At the present time, in the
majority of places in Turkmeniya, except remote sections of the
sands, the corsac fox is common (Sapozhenkov, 1960; V.G.
Heptner). In Tadzhikistan, corsac fox is rare in the flat southern
and southwestern regions. In the period from 1951 to 1959, only 11
(1952) to 63 (1959) skins were tanned per year. As mentioned
above, corsac fox is absent in montane Tadzhikistan.

In Uzbekistan, the corsac fox occupies in all of the plains and
foothill desert and semidesert sections, and is common in some
places. In the 1924/25 season, 8,039 skins were prepared in the
republic and in 1925/26—5,442 (Kogan, 1931). In recent years, the
number of animals everywhere decreased (Ishunin and Lustin, 1959),
probably in connection with the intensive development of new lands,
but in a series of districts, it is common at the present time. Thus,
from 1956 to 1959, in Kashkadar’insk district, from 580 to 1,244
corsac foxes were caught per year, in Bukhara—from 395 to 546,
in Samarkand—from 288 to 474, and in the Tashkent—from 208 to
336. In the Fergana valley, they are obtained singly from 1956—

284

412

1959, 1-12 skins. The corsac fox is rare in Korezmsk (16—88 skins
per year) and Surkhandar’insk districts (68—94 skins per year).

In Kirghiziya, in the 60’s of the previous century (N.A.
Severtsov, 1873) the corsac fox was common not only in the Chuisk
and Talassk valleys, but also in high mountane regions (Susamyrsk
Plateau, Dzhumgal, the Lake Sonkul’ depression, on the Aksai and
in the lower reaches of the Chirchik, Arys, and Keles rivers).
However this information is doubtful. In the last decade, the corsac
fox was not found permanently in any place in Kirghiziya (Kuznetsov,
1948). In the severe winters of 1948/1949 and 1950/1951, these
animals appeared in the Chuisk valley and the foothills adjacent to
it, and became fairly common there. In 1956—1959, Kirghiziya yielded
131-137 skins of this animal.

In southeastern Kazakhstan (southern Pri-Balkhash’, Chu-Iliisk
mountains, foothill regions of the Zailiisk and Dzhungarsk Alatau,
Alakul’sk depression), the corsac fox was extremely rare and was
caught singly until the 50’s of the present century. In many regions,
it was completely unknown. Starting from 1951, populations of the
animals here started to grow rapidly and in recent years in Alma-
Atinsk and Dzhambulsk districts, up to 1000—1300 skins began to be
taken annually.

In southern Pri-Baikal the corsac is “very rare” (Selenga valley;
Fetisov, 1949) and does not appear every year (Surmach, 1957). In
southeastern Trans-Baikaliya in the middle of the last century, the
corsac fox was very common in the montane steppes (Maak, 1859).
“Along the Argun’ and Onon they are a multitude” (Cherkasov,
1884). Later, in connection with the ploughing of the steppes, num-
bers of these animals strongly decreased in eastern Siberia and in
the 20’s of the present century only 341 skins were tanned in 1925/
26 and in 1926/27, 333 skins. In the 30’s, the number of corsac
foxes here again grew, and up to 1,400 corsac began to be caught
in a year (Podarevskii, 1936). In the last two decades, the number
of the corsac foxes in Trans-Baikaliya has again decreased (see
“Population”).

Therefore, within the boundaries of USSR, the corsac fox is
most numerous in dry steppes and semideserts (central Kazakhstan).
In deserts of the northern and southern types (southern Kazakhstan,
Turkmeniya, Uzbekistan) this carnivore is less numerous, but in
places it is common and even abundant.

413 .

In the 1 8th century, in the territory of contemporary Kazakhstan,
corsac was much more numerous than now. At that time} Kazakhs
sent from 40 to 50 thousand skins every year to Orenburg and
Troitsk fortresses (Rychkov, 1762). “Innumera copia” (innumer-
able multitude)}—thus was described the number of corsac foxes in
Kazakhstan at the end of the 18th century and beginning of the
19th by G. Fisher (1814). Later, as a result of intensive harvest and
especially ploughing of the steppes, the populations of corsac fox
started to decrease. They declined especially strongly after 1954 in
connection with massive ploughing of virgin lands.

The corsac fox is numerous in the Mongolian People’s Repub-
lic. In individual years, up to 18 (1928) and even 24 thousand skins
(1931) were obtained there. About 90% of this quantity was ob-
tained from the eastern plain-steppe and semidesert regions
(Bannikov, 1954). In the province of Xinjiang (China), in Iran and
in Afghanistan, the corsac fox is rare. In northern Afghanistan,
however, it 15 quite common (V.G. Heptner).

Habitat. In the Pri-Azov the corsac fox is restricted to the
weakly undulating plains of the watersheds which are occupied by
Pri-Black Sea herbaceous-fescue-feather grass and fescue-feather
grass steppes, as in the southern chernozem. Here are innumerable
ravines and gullies, at the heads of which the soils are salinized and
have semidesert vegetation, covered mainly by fescue, meadow
grass (Poa bulbosa) and desert-steppe shrubs: prostrate summer
cypress (Kochia prostrata) and various species of sagebrush; also
thickets of steppe shrubs—blackthorn (Prunus stepposa), spiraea,
Russian peashrub (Caragana frutex) and others. These steppes
were formerly inhabited by bobac marmots [M. bobac]. At the
present time all of these steppes are almost completely destroyed,
in connection with which the corsac fox lives there on the solon-
chak [areas of saline soil] near the seashore (Korsak-mogila)
[Corsac-grave], in elevated areas; and along southern slopes of
gullies as well as in pastures.

In the western half of the Cis-Caucasus, this carnivore lives in
steppes very similar to those already described, while in the Pri-
Caspian and the lower Volga, the corsac fox inhabits sagebrush-turf-
grass steppes and semishrub sagebrush and Russian thistle desert
(southern semidesert), and is also encountered on compacted sahds
(Nogaisk steppe). In the Volga-Ural interfluve, the corsac fox in-
habits the steppes, semideserts and deserts, but it is more common

285

414

in hilly semideserts with a low and scattered stands of grass; along
the middle course of the Ural river, on the Obschii Syrt and on the
upper Chagan river, it lives in the hilly steppes (Zarudnyi, 1894) and
chiefly inhabits the slopes of hills scattered across the level feather
grass-sagebrush semidesert (Kuznetsov, 1928). In the Volga-Ural
sands, it is rarely met with and, apparently, is not permanent.

During years of increase, it inhabits even the forest-steppe of
western Siberia beyond the Ural mountains, with its flat relief where
the low and narrow ridges (“crests”) 2 to 4 m high, rarely 6 to
10 m, alternate with the depressions occupied by lakes, rush-cov-
ered floodlands, small birch forest—kolka [“pins”}—and
solonchak. In the forest-steppe, the corsac fox mainly inhabits
solonetz [alkaline] soils around large salt lakes with poor vegetation
and occasionally the crests if they contain significant sections of
virgin soil with low fescue steppe. It hunts mainly around water
bodies with open shores, on solonchak, in depressions and along
crests where little snow occurs in winter. It flees into kolka and
rush-covered floodlands only during periods of ice crusts, where it
searches for white hares [Lepus timidus] and willow ptarmigan
[Lagopus lagopus] caught in snares (Sludskii, 1930).

South of the forest-steppe, from the left bank of the Volga and
to the Ob’ [river], the corsac fox lives in herbaceous-fescue-feather
grass steppes on ordinary and southern chernozem (Kulundinsk
steppe), and also in turf-grass dry steppes lying more to the south.
In the typical steppes, the corsac fox settles in the burrows of
marmot and ground squirrels, located on the gentle slopes of
elevated areas not far from the edges of solonchak, and in sum-
mer it hunts mainly in places with scarce vegetation. There, where
the steppes are ploughed, the corsac fox survives only in pastures
and near bodies of waters; it is often obliged to live near villages.

In winter, the corsac fox chooses places where the snow is
blown clear by wind or is light, but it also appears in thickets of
reeds near bodies of water. Thus, in the very snowy winter of
1940/41, in Severo-Kazakhstansk district, corsac foxes were found
in considerable numbers on the steep slopes of the great Lake
Chagla depression, where there was little snow and it could prey
on voles and mice. Driven by hunger, the corsac foxes were cap-
tured in cut reeds growing around the lake, under which there was
little snow and where they hunted the water voles [Arvicola
terrestris] which were very numerous there.

415

In Kazakhstan, the corsac fox is especially numerous in desert
steppes between the Ural and Emboi, along the Turgai and south
of Lake Kurgal’dzhin and the Nura river. There is undulating or
hilly relief with narrow river valleys and quite expansive depres-
sions with fresh and salt lakes. In these regions, the corsac fox
occupies gentle sides of valleys and slopes and is usually absent in
melkosopochnik [region of small mounds or hills], occurring only
around the base of the sopok [mounds]. Thus, southeast of Lake
Kurgal’dzhin, near the Kulanutmes river, burrows were distributed
on the upper parts of the slope—the watershed between the
Kulanutmes and a deep narrow valley. From the river to the sum-
mit of the slope was 2—3 km. The gentle declination of the slope
was occupied by fescue-feather grass steppe with 30-50% of the
soil covered with vegetation. In the upper part of the slope, old
burrows of marmots and ground squirrels were encountered. Corsac
foxes also used them as dens.

The building of a burrow by corsac foxes on elevated areas (ex-
cluding melkosopochnik) is explained by the high flooding of the steppe
rivers; the water level of which rises 10—14 т. Building their burrows
in the higher part of knolls, the corsac fox in summer hunts along their
slopes, and rarely appears at the bottom of ravines, or at the river or
lake, but it usually avoids entering hilly country. In autumn and the
beginning of winter, in the northern semidesert (Lake Kurgal’dzhin), it
restricts itself to flat areas, overgrown with sagebrush and Russian
thistle (B.D. Abaturov).

After abundant snowfall, when the depth of snow cover in
areas with tall vegetation reaches 20 to 40 cm, and it becomes
more densely packed, while in areas with low vegetation cover (in
sagebrush, sagebrush-fescue and sagebrush-saltbush [Atriplex cana]
associations), snow cover is shallow; the corsac fox prefers such
sections with little snow or places with very packed snow. There,
the winter burrows are also found, located along more elevated
areas.

Farther to the south, the corsac fox lives in true deserts of the
northern type, inhabiting the entire Betpak-Dala. In summer, it is
rarely encountered here. For nine years, on an itinerary extending
for a total of ten thousand kilometers, only four burrows with litters
were found. In Betpak-Dala, the corsac fox settles on the summits
of knolls covered with small sagebrush and usually 200—300 т from
a spring with water. In the last decade, corsac foxes became

286

416

common in the foothills of Zailiisk Alatau, where they inhabit hilly
clayey deserts and semideserts, not long ago inhabited by yellow
ground squirrels [Spermophilus fulvus], in the burrows of which
the foxes construct their lairs. Here, the corsac fox finds favorable
conditions, since food is abundant, snow in winter is not deep, and
is often trampled down by numerous cattle herds grazing there.
The corsac fox inhabits the ephemeral deserts of Turkmeniya,
Tadzhikistan and Uzbekistan. Along the lower Amu-Dar’ya, it oc-
cupies the old valleys and deltas, but everywhere it is quite rare
(Kostin, 1956). In clayey deserts of southwestern Turkmentya, it is
met with everywhere, and is not rare in compact sands. In south-
eastern Turkmentya, the corsac fox is found in the clayey deserts
and semideserts and along the edges of compact sands. It is com-
mon in the southern margins of the Karakum; it often remains near
wells. There, its burrows are found on the slopes of sandy hillocks
inhabited by the great gerbil [Rhombomys opimus]. In Tadzhikistan
and Uzbekistan, it inhabits desert areas on the plain and in the

Fig. 109. Burrow of corsac fox. Betpak-Dala desert. Kazakhstan. 16 June,
1956. Photograph Бу A.A. Sludskil.

287

417

foothills, old river valleys and the small sandy massifs. It is absent
in tugai [riparian] forests (Chernyshev, 1958).

In southern Pri-Baikal and southeastern Trans-Baikal, the corsac
fox inhabits hilly steppes, well-supplied with mounds having flat
summits 50-100 m high which are irregularly scattered or extending
as low chains. The slopes of the sopok are gentle, and pass gradu-
ally into wide depressions. In places on the hills and in the steppe,
there are rocky exposures and outcrops. Among the rocks and near
them, forbs and shrubs grow vigorously which are rarely encoun-
tered in the ravines. On large areas, the grassy steppes stretch over
sandy and sandy-gravelly soil, mainly wild calla* cattail and wild
rye-cattail steppes. In the steppes, the grassy and sagebrush areas
form a complicated mosaic. The region is characterized by low
temperatures in winter (to — 50°C), shallow snow cover—on aver-
age only 10 cm—and permafrost located at a depth of 2.0—2.6 т.
In Trans-Baikaliya, the corsac fox inhabit the slopes of valleys
well-supplied with tarbagan [Marmota sibirica] burrows which
they occupy for their shelters.

In the MNR [Mongolian People’s Republic], the typical habitat
is more or less level or slightly undulating dry steppes with small,
shallow valleys with a cover of feather grass, feather grass-bistort
or grass-Russian thistle. This carnivore is rarer in herbaceous steppes
of the northern type, and still more rarely encountered in montane
steppes, whence it penetrates along the river valleys. It avoids
mountainous relief with cliffs and rock slides as well as true shrub
deserts.

Accordingly, throughout the entire range of the corsac fox, the
main habitats are the semideserts and the dry steppes resembling
them. In grassy steppes, it chooses areas with scattered, low grass
and shrub cover. Everywhere, it avoids dense thickets and prefers
the plains. The corsac fox is absent in areas with loose snow cover
more than 15 cm in depth, but there, where snow is compact or
blown away by wind, it may exist even when the average depth is
30 to 50 cm deep.

Food. The corsac fox is a carnivore, adapted to hunt small and
moderate-size lagomorphs and rodents, including young marmots
and hares. In summer time, small birds, reptiles and insects also
occupy a considerable place in its diet. In distinction from other

*The Russian word “zmeevik’ is used for 5 different genera, including
Polygonum, Calla and “tyrsa” is defined as Typha—Sci. Ed.

288

418

species of the family of our fauna, it rarely feeds on plant food. In
case of insufficient prey, it eats carrion and refuse, often poorly-
digestible.

In connection with the wide distribution of the corsac fox, its food
list varies, but everywhere rodents predominate. At the northern
limits of the range, in the forest-steppe of western Siberia (Barabinsk
steppe), this carnivore in summer preys on narrow-skulled voles
[Microtus gregalis| or Alticola strelzovi and sagebrush voles
[Lagurus lagurus]; rarely does its prey extend to water voles
[Arvicola terrestris], red-cheeked ground squirrels [Spermophilus
erythrogenys], great jerboas [Allactaga jaculus*], common ham-
sters [Cricetus cricetus], Dzhungarian hamsters [Cricetulus
sungorus], larks, charadriiformes and ducks. In winter, it preys
mainly on mouse-like rodents, but occasionally attacks white hares,
willow ptarmigan, gray partridge [Perdix perdix], black larks
[Melanocorypha yeltoniensis| and snow buntings [Plectrophenax
nivalis]. Near villages, it feeds on carrion and refuse; several in-
dividuals gathering around a large carcass (Sludskui, 1930). In the
steppes of Kazakhstan, a basic role in the food is also played by
mouse-like rodents (99.4%), among them the sagebrush vole (37.9%).
Apparently, this rodent is more often eaten, since of the numerous
remains of unidentified voles found in the feces a significant part
also (48.5%) belong to it (Table 32). In this region, birds are also
important in the food of corsac fox (35.5%), apparently wounded ones.

In the steppes in winter, the corsac fox feeds on mouse-like
rodents, but when they are scarce or not obtainable, it turns to other
species of prey. Thus, in the heavy snow winter of 1940/41, in
Severo-Kazakhstansk district, at Lake Chagla during a time of fro-
zen snow-crust, corsac foxes hunted water voles in cut reeds,
digging up their runways in the thawed soil, and also gray partridge
which spent the night in shrubbery near the lake. These carnivores
also remained near cattle burial grounds and villages, where they
fed on carrion and refuse.

In 890 feces of corsac fox collected in May—June 1947 and
1948, near Tselinograd, remains of rodents were found in 95.4% of
all samples investigated, corresponding to: birds in 17.0%, insects in
22.1%, reptiles in 0.8%, plants remains in 2.1% and indigestible
refuse (rags, ropes, etc.) in 0.4% (A.F. Chirkova). Among mouse-
like rodents, the main mass consisted of sagebrush voles, while

*Now called А. major—Sci. Ed.

419

287 Table 32. Results of feces analysis of corsac fox inhabiting the Kazakhstan
steppes (Severno-Kazakhstansk district, Sovetsk region, near Lake Chagla,
July, 1947; from burrow with litters; 169 samples; frequency of occurrence

in %)
Food type %

Mammals 99.4
Rodents 99.4
Various voles 99.3
Including:

Sagebrush vole [Lagurus lagurus]| 37.9

Common vole [Microtus arvalis] 0.6

Narrow-skulled vole [Microtus gregalis] Ta

Root vole [Microtus oeconomus | 4.2

Unidentified voles 48.5

House mouse [Mus musculus] 0.6
Birds 35.5
Ducks 14.8
Insects al
Beetles 1-1

common, root and narrow-skulled voles, mole-vole [E/lobius] and
wood mice [Apodemus] were rarer. The remains of white hare,
red-cheeked ground squirrel, jerboa, shrews—brown-toothed [Sorex]
and white-toothed [Crocidura}—were found on isolated occasions.

In the semideserts and dry steppes of central Kazakhastan in
the spring—summer period, corsac foxes hunted mainly intensively
reproducing sagebrush voles (encountered in 96.1% to 98.7% of all
samples examined). Other voles, rats, hares, steppe pika [Ochotona
pusilla], hamsters [Cricetus] and dwarf hamsters [Cricetulus] were
rarely met with in the feces and clearly played no notable role in
its diet. At the same time, the role of ground squirrels and jerboas
here was increased. Thus, e.g. at Karsakpai, the remains of yellow
ground squirrels were found in 13.3% of the samples, while in the
Atbasarsk region of Tselinograd district, in 7.1%, although there this
ground squirrel lives at the northern limit of its range. In the latter
region, the remains of intermediate ground squirrels [Spermophilus
intermedius|* were found in 7.2% of the excrements, while those
of jerboas—in 14.2% (Kh. Kydyrbaev). In the Basaga region, jer-
boas were found in 10.7% of the samples (see Table 33).

*Synonym, 5. brevicauda; usually considered a subspecies of
S. erythrogenys—Sci. Ed.

420

289 Table 33. Results of feces analysis of corsac fox inhabiting the semidesert
of central Kazakhstan (in % of number of samples)

Region, season Northern Southern semidesert—
and year, number — semidesert near Karsakpai,

of samples Kulantumes Kazakhstan upland,
river Basaga’

Summer, Summer, Spring, Summer, Summer,
1950 1951 1950 1953 1954
417 1518 45 103

Food type

Mammals 99.8 99.4 100.0 100.0 100.0

Lagomorphs and rodents 99.8 99.3 100.0 100.0 100.0

Steppe pika [Ochotona pusilla} — 0.1 2.2 4.0 3.8

Mongolian pika [O. pallasi] — — — 2.0 eS

White hare [Lepus timidus] 0.2 0.4 — — —

Yellow ground squirrel — — 13.3 — —
[Spermophilus fulvus |

Ground squirrel [S. intermedius] 0.5 0.1 — 5.8 —

Little ground squirrel — — 22 — —
[S. pygmaeus]

Ground squirrel (unidentified) 0.5 0:2 — 13.8 —

Steppe mouse [Sicista subtilis] — 0.3

Great jerboa [Allactaga jaculus] — 0.5 — — —

Small five-toed jerboa — — 2.2 — —
[A. elater]|

Thick-tailed three-toed jerboa — — — 2.0 —
[Stylodipus telum]

Jerboas (unidentified) 1.0 4.3 — 10.7 0.8

Common hamster — 0.1 — — —
[Cricetus cricetus |

Eversmann’s hamster 0.7 0.1 — — —
[Cricetulus eversmanni|

Gray hamster [C. migratorius] — — — 3.0 т

Sagebrush vole 98.7 96.6 75.6 96.1 97.0
[Lagurus lagurus|

Mole-vole [Ellobius] 2.9 0.1 — — —

Water vole [Arvicola terrestris] © — 0.1 — — =

Common vole 0.2 — 8.9 — —
[Microtus агуа $]

Narrow-skulled vole — — — 9:7 12.1
[М. gregalis|

House mouse [Mus musculus | — 0:7 — — aS

Striped field-mouse 0.2 — — т =
[Apodemus agrarius |

Wood-mouse [A. sylvaticus] === 0.4 = — =—=

Small rodents (unidentified) 0.1 0.3 — — =

Carnivores 0.5 0.5 ЗЕ oe ue
(сота.)

421

(Table 33—contd.)

Region, season Northern Southern semidesert—
and year, number — semidesert near Karsakpai,

of samples Kulantumes Kazakhstan upland,
river Basaga’

Summer, Summer, Spring, Summer, Summer,
1950 1951 1950 1953 1954
417 1518 45 103 132

Food type

Weasel [Mustela nivalis] 0.5 0.3 — — —
Corsac fox (wool) — 0.2 = — —
Birds 18.7 16.7 — 5.8 6.8
Ruddy Shelduck 0.2 — — =e —
Ducks (unidentified) 1.2 — — — —
Large birds 4.6 0.7 — — —
White-winged lark 2.4 0.9 — — —
Small birds (unidentified) 10.6 16.0 -— -— —
Duck eggs 0.2 0.1 — — —
Reptiles fey 8.1 — 2.0 les
Sand lizard 0.1 1.3 -— — —
Lizards (unidentified) 0.5 | — 2.0 | 55
Patterned rat-snake — 0.1 — — —
Mamushi — 0.1 — — —
Snakes (unidentified) 0.2 0.5 —- — —
Insects 22.0 17.4 — 15.8 4.5
Locust — т — — —
Beetles 22.0 16.3 — — —
Eggs of praying mantis — 0.1 — — --
Plants (grasses) 0.2 0.1 — — —

Carrion — 0.1 — — —

7According to the data of G. Shubin.

In places where the tarbagan marmot is still numerous, it appar-
ently plays a considerable role in the summer diet of corsac foxes,
since in Trans-Baikaliya, corsac foxes catch young bobaks and
feed on the dead bodies of adult animals (see below).

In the semidesert, the role of reptiles among the corsac fox’s
food is negligible (1.5—3.1%). The same may be said about insects
in years of rodent abundance (see Table 33). In a time of depres-
sion of mouse-like rodent populations (1960), in the region of Lake
Kurgal’dzhin, the feces of corsac fox consisted, to a considerable
degree, of remains of grasshoppers (B.A. Abaturov). Birds have a
more basic importance in the food of the corsac fox in semideserts
(5.8-18.7% occurrence). In summer, this carnivore almost never
eats carrion, but of plants it consumes small quantities of grass
leaves.

290

422

In winter, with sagebrush voles abundant, and obtainable, the
corsac fox continues to hunt this rodent principally, and, for ex-
ample, in the region of Lake Kurgal’dzhin in the winter of 1959/60,
all of its feces consisted entirely of the remains of mouse-like
rodents. Moreover, in river floodplains, they successfully attack
white hares; hunt black larks hiding in the snow at night from bad
weather. Holes of corsac fox were found in early spring which
contained shells of bird eggs buried by them in the ground. In the
winter of 1960/61 in the region of Lake Kurgal’dzhin, when sage-
brush voles were very scarce, corsac foxes restricted themselves
to harvested fields and roads. In fields, they hunted harvest mice
[Micromys minutus], house mice and common voles, and also gath-
ered the seeds of cereal crops. In winter, feces were, mainly,
mouse-like rodents, but contained also remains of seeds of corn
[maize], oats and barley. The hungry animals dug out from under
the snow old boots, soles, rags, ropes and gnawed them (B.A.
Abaturov). There, they also picked up small fish discarded by fish-
ermen. In Tselinograd district in the winter of 1940/41, which
distinguished itself by its abundant snow and frozen crust, corsac
foxes destroyed white and common hares so intensely that their
number became considerably reduced. An analogous picture was
also observed there in the winter of 1953/54. After a thaw and the
formation of a strong ice crust, corsac foxes began to starve and
appeared in great numbers in the river floodplains and clearings,
where they intensively hunted white hares or remained near vil-
lages where they fed on refuse. Feeding of corsac foxes in winter,
a time of famine for them, on refuse near populated areas, and
even the outskirts of such large cities as Pavlodar and Karaganda,
is a common phenomenon.

In deserts of the northern type, extending in Kazakhstan from
the northern Pri-Caspian to the Alakul’sk depression, rodents con-
tinue to predominate in the summer food of corsac foxes (87.3—100%
of all occurrences) (see Tables 34 and 35), but the importance of
different species of animals changes greatly. Mouse-like rodents,
especially the sagebrush vole [Lagurus] stand as secondary* prey.
Only in the foothills of the Zailiisk Alatau and in a few other similar
places, was the social vole [Microtus socialis| encountered in
70.8% of occurrences, and in the foothills of Bulattau (northeastern
Betpak-Dala)—in 38.4% of samples studied. 3

*From context, probably should read “ргитагу”—$с1. Ed.

2911

423

Table 34. Results of feces analysis of corsac fox inhabiting northern
type clayey deserts in Kazakhstan (in % of sample number)

Region, year, num- Pri-Caspian Foothill, Alakul’

ber of samples lowland, Zailiisk depression,

Makat Alatau, Uch-Aral
Samsy

May May—June April-June,
1934 1954-1955 1954
7988 2099 131%

Food type

Mammals — 99.0 100.0
Lagomorphs and rodents — 99.0 100.0
Hares [Lepus sp.]* 1.0 — —
Yellow ground squirrel [S. fulvus] — 40.2 —
Little ground squirrel [5. pygmaeus] 12.0 — —-
Great jerboa [Allactaga jaculus| 0.6 0.5 —
Earth hare [А. jaculus]** 1.1 — —
Small five-toed jerboa [A. elater] — 1.0 —
Fat-tailed jerboa [Pygerytmus platyurus] 0.2 — —
Thick-tailed three-toed jerboa 4.0 — —
[Stylodipus telum]
Jerboas (unidentified) 19.1 -— 5.3
Migratory (gray) hamster — — 19.1
[Cricetulus migratorius]
Eversmann’s hamster [C. eversmanni] 1.6 — —
Tamarisk gerbil [Meriones — 11.5 2.3
tamariscinus]
Mid-day gerbil [M. meridianus] 0.6 — —
Gerbils (unidentified) 3.5 — —
Mole-vole [Ellobius] — 6.0 —
Muskrat [Ondatra zibethicus] — — 13.0
Social vole [Microtus socialis] 8.2 70.8 5.3
Root vole [М. oeconomus| — — 7.3
Voles (unidentified) —- —- 22.9
House mouse [Mus musculus | — 6.0 15
Mouse-like rodents 22.5 — 29.6
Rodents (unidentified) 7.0 — —
Carnivores — 0.5 —
Steppe polecat [Mustela eversmanni] — 0.5 —
Insectivores 1.5 — =
Long-eared hedgehog 15 — —

[Hemiechinus auritus]
(contd.)

*In the Russian original, zayats-rusak, which is the name applied to L.
europaeus. This is evidently a /apsus, and L. tolai is meant—Sci. Ed.

**In the Russian original, zemlyanoi zaichik, another common name for the
great jerboa. Probably a /apsus—Sci. Ed.

424

(Table 34—contd.)

Region, year, num- Pri-Caspian Foothill, Alakul’

ber of samples lowland, Zailiisk depression,

Makat Alatau, Uch-Aral
Samsy

May May—June April-June,
1934 1954-1955 1954
2099 130%

Food type

Birds 10.5 3.0 3.8
Gray partridge [Perdix perdix] 0.3 — —
Merlin [Falco columbarius| 0.1 — —
Kestrel [Falco sp.] 0.1

Gray lark [Calandrella pispoletta| 0.2 — —
Passiformes (unidentified) —- 3.0 2.3
Birds (unidentified) 9.8 — 1.5
Reptiles 1.4 0.5 43.5
Lizards — 0.5 30.5
Snakes — — 13.0
Insects 1.6 16.0 —
Locust — — 1:5
Mole crickets — — 38.9
Beetles — 16.0 297
Insects (unidentified) — — 45.0
Plants (grass leaves, seeds of 1.5 6.0 6.9

barley and wheat).
Carrion (of sheep, camels) Del — —

8Kolosov (1935).
°*Kh. Kydyrbaev.
10K husainov (1959).

In the food of desert corsac foxes, jerboas play an important
role. In the Pri-Caspian lowlands, they were noted in 25%, while in
the clayey-gravelly Betpak-Dala—in 30.2—77.8% of examined samples.
Rodents of medium size occupy second place. In the Pri-Caspian
lowland, these are the little ground squirrel [S. pygmaeus] (12.0%
occurrence), in the northwestern part of the Betpak-Dala—the in-
termediate ground squirrel [5. intermedius] (43.7—61.9% occurrence)
and the steppe pika [O. pusilla] (53.8% occurrence); lastly, in the
foothills of the Zailiisk Alatau—the yellow ground squirrel [5. fulvus]
(40.2% occurrence) (see Tables 34 and 35). In the southern part
of Betpak-Dala, the great gerbil [R. opimus] was noted in 92.0%
of feces examined, and on the edge of Taukum (southern Pri-
Balkhash)—in 40%. In the latter region, jerboas (40%), social voles

292

425

Table 35. Results of feces analysis of corsac fox inhabiting desert of
the northern type—Betpak-Dala (in % of sample number)

Region, year Southern Central Northeastern
number of samples (Koktal) (Sorbulak) (Bulattau)

June May May May
1955. 1954, 1956, Т95Ё
112" 103 63 13

Food type

Mammals — 100 100 100
Lagomonrphs and rodents — 96.1 87.3 9253
Steppe pika [Ochotona pusilla] — 54.4 —- —
Mongolian pika [O. pallasi] — — — 53.8
Tolai hare [Lepus tolai] 4.5 — —- —
Intermediate ground squirrel 11.6 43.7 61.9 7.7
[5. intermedius |
Jerboas (unidentified) 42.0 77 30.2 23.0
Migratory (gray) hamster 7 1.0 — —
Great gerbil [R. opimus] 92.0 — — —-
Voles (unidentified) — 1.0 1.6 38.4
Carnivores = === — Usd)
Weasel [Mustela nivalis | — — — Thal
Ungulates — 2.9 11.1 ---
Saiga [Saiga tatarica] — 29 7.9 —
Goitered gazelle [Gazella — — 3.2 —
subgutterosa|
Insectivores 1.8 1.9 3:2 —
Long-eared hedgehog [Hemiechinus 1.8 1.9 3:2 —
auritus |
Birds (passiformes) 2.7 12.6 31.7 Teh
Reptiles Ths 1:9 15.9 1
Lizards 6.2 1.9 7.9 —
Snakes 0.9 — 7.9 ИЯ
Insects 34.0 19.4 36.5 15.4

Beetles 34.0 19.4 36.5 15.4

(20%) and insects (beetles) (40%) of feces studied are of greatest
importance in food of the corsac fox.

Near a burrow with a litter (14 May 1952, at the edge of
Taukum desert), remains of black-bellied sandgrouse and steppe
larks were found, and at a family burrow in the Ili’ delta (May
1961), remains of house mice, narrow-skulled voles [М. gregalis],
muskrats and Semirechensk pheasants were found. The presence
of remains of saiga and goitered gazelle in corsac fox feces in

ИМ.1. Ismagilov.

293

426

spring must be attributed to carrion feeding. In some deserts birds,
mainly passeriformes, sometimes occupy an essential place for the
described predator (up to 31.7% occurrence) (see Table 35). Corsac
fox attacks even such large birds as graylag goose [Anser anser].
At the end of July 1936, on Lake Telikul’, was observed how it
concealed itself at midday from adult birds sitting on the shore. It
successfully attacked young wild geese and ducks on Barsa-Kel’mes
Island (Aral Sea; Sludskii, 1939).

The corsac fox readily feeds on lizards, snakes (eastern snake,
chicken and rat snakes) and young steppe [Horsfield’s] terrapins
especially in years with low numbers of rodents. In the Alakul’
depression, in spring and summer 1954, when the number of ro-
dents was low, 43.5% of corsac feces near burrows with litters
contained remains of reptiles, mainly lizards (30.5%). Insects are
regularly eaten by corsac foxes in the desert and in considerable
quantities, but usually do not play an important role in the diet.
During insufficiency of rodents, they occupied a prominent place in
the nutrition of a corsac fox family in Alakul’ depression in 1954
(see Table 34). Mole crickets (38.9% occurrence) and beetles are
particularly often eaten. Plant remains are rarely found. Usually,
these are grass leaves, and in the foothills of the Zailiisk Alatau, the
animals in spring gleaned seeds of wheat and barley remaining on
the soil surface after sowing.

When it turns cold and the snow falls in the desert and semi-
desert zone, the corsac fox is deprived of many foods: rodents
entering into winter torper, migratory birds, reptiles and insects. The
remaining voles, pikas and gerbils which retain an active mode of
life exhibit reduced surface activity or shift to a subnival mode of
life. Therefore, the majority of corsac foxes leave summer habitats
and migrate in search of food to the river valleys (Ural, Emba, Syr-
Dar’ya, Chu, Ili), and to sand [deserts] (Pri-Aral Karakam, Pri-Chu
Muyunkum, Sary-Ishikotrau), and sometimes northward into the dry
steppes. In winter both migratory and sedentary animals feed mainly
on the remains of saigas and goitered gazelles preyed on by wolves,
and on carrion. In searching for carrion and refuse, corsac foxes
move at that time nearer to populated places and roadbeds of
railway lines. Feeding on the remains of the wolf’s prey, corsac
foxes become their true commensals. For instance, in the Ural-
Kushum interfluve, in the winter of 1953/54, in December—January,
corsac foxes fed mainly on the carcasses of male saigas killed by

427

wolves. In 10 animals, caught in the Betpak-Dala desert from
November 27 to December 21, the remains of saigas were found
in 70% of the stomachs. In the middle of December, 1955, in the
central part of Betpak-Dala, several corsac foxes were found at
each saiga carcass.

Food of the corsac fox in southern (ephemeral) deserts has not
been studied. Judging by the faunal composition of these deserts,
one may suppose that in summer it feeds there, apparently, on great
and red-tailed [Libyan] gerbils, yellow ground squirrel, various
species of jerboas and birds, reptiles and insects, and in winter, on
gerbils and rarely on jerboas and birds.

In the steppes of southeastern Trans-Baikaliya summer food of
the corsac fox consists mainly of rodents (99.2—100% occurrence)
(see Table 36). It quite often feeds on tarbagan marmots (18.8—
71.0% occurrence) and Daurian pika (29.2 to 62.8% occurrence).
Voles and hamsters stand in second place in its food, then birds,
and lastly—tolai hare, Daurian ground squirrel, jerboa [Allactaga
sibirica], insectivorous and carnivorous mammals, and insects. Char-
acteristically, in Trans-Baikaliya, reptiles, which are rare there, are
lacking in the food of corsac fox. It more often feeds on plant than,
for example, in the steppes of Kazakhstan.

The corsac fox, attacking a tarbagan marmot, causes relatively
little harm, except to the young. These rodents do not especially
fear corsac foxes and rarely conceal themselves in burrows on
their approach. They only interrupt feeding, sit “like a stump”, and
call. Not rarely one may observe how an adult tarbagan marmot
attacks a corsac fox, jumping out from its burrow or drawing nearer
and nearer, and drives it away from the mound. Tarbagan marmots
a year old or more, on being kept in a cage with an adult corsac
fox, remained alive during the course of several days, notwithstand-
ing that the carnivores were not fed during this time. Under similar
conditions, young tarbagan marmots (several months in age) quickly
became victims of the corsac foxes (Brom, 1952). Remains of 23
tarbagan marmots, found near and in the burrow of a corsac fox
consisted of: juveniles (mendel)—4, animals in their second year
(kotel)—16; and adults—3 (Nasimovich, 1951). It is believed that the
corsac fox can kill only young tarbagan marmots, but among the
adults, it takes solely dead ones (Nasimovich, 1951; Brom, 1952).
Sometimes marmot families with young live next to a corsac fox
litter and the carnivores do not touch them.

428

294 Table 36. Food of corsac foxes in the steppes of southeastern Trans-
Baikal based on analysis of samples from contents of their stomachs.
and feces (in % of sample numbers)

Season of Summer Winter

year, material

52 feces feces
stomachs'’® 56 500 sto- 323
Food type samples'* sam- machs" samples
ples'?
Mammals 100.00 100.00 99.4 52.6 —
Lagomorphs and rodents 100.00 100.00 99.2 39.4 94.2
Tolai hare [Lepus tolai] — 5 — 7.9
Daurian pika [Ochotona 32.6 29:2 62.8 — 5
ааитса]
Daurian ground squirrel 3.8 Tcl! 1.0 — —
[Spermophilus ааитси$]
Tarbagan marmot [Marmota 71.0 60.0 18.8 557 50.8
sibirica]
Jumping jerboa 1.9 6.1 2.0 2.6 0.3
[Allactaga sibirica saltator]
Daurian [striped] hamster — — — 10.5 2.5
[Cricetulus barabensis]
Dzhungarian hamster 14 1.5 — 2.6 2.8
[Phodopus sungorus}
Hamsters (unidentified) Tl 12.3 0.6 — 6.8
Trans-Baikal {5оКог. [ Муозра!ах] — — — 2.6 —
Michnoi vole [Microtus michnoi| — — — 5:2 —
Narrow-skulled vole tl 9.2 — 2.6 3.7
[М. gregalis]
Brandt’s vole [М. brandti] 9.6 6.1 — 7.9 5.6
Voles (unidentified) 1.9 12.3 13.0 21.0 127
House mouse [Mus musculus] — 5 — —- 0.3
Murids (unidentified) 5.8 3.0 16.8 —- —
Chiroptera — ].5 — — —
Insectivores 1.9 3.0 — 7.9 1.2
Daurian hedgehog [Mesechinus — 3.0 — 2.6 1:2
dauuricus}
Red-toothed shrews [Sorex sp.] 1.9 — — 5.2 —
Carnivores el 1.5 0.2 5.2 №2
Steppe polecat [Mustela — — — 5-2 —
eversmanni|
Solongoi [M. altaica] — — 0.2 —
Corsac fox 7.7 les) — — 1.2
Ungulates 3.8 15.6 0.2 2.6 3.4
Mongolian gazelle [Procapra — 15.6 — — 3.4
вийего5а]

(contd.)

429

(Table 36—contd.)

Season of Summer Winter
yeatmatenlalne ojo anatase lis a

hag hg eae a ae
stomachs’? 56 500 sto- 323
Food type samples’? зат- machs'* samples
ples’?
Roe deer [Capreolus capreolus] — — — 2.6 —
Carrion (domestic animals) 3.8 27.6 0.2 23.7 32.7
Birds Ted 153 33.4 26.3 13.0
Daurian partridge [Perdix — — —- 7.9 —-
dauurica|
Mongolian lark — — — 2.6 —
[Melanocorypha топгойса]
Small birds — — — 10.5 —
Insects 55:7 27.6 9 2:6 3.4
Plants 32.7 37.0 1.8 10.5 30.6
Refuse (pieces of twine, rags, — 3.0 —- 59.9 9.3
dressed skin, etc.)
Soil, pebbles 63.4 63.0 — — 133

Brom (1952).
'83Nasimovich (1951).
“4Skalon (1936).

The negligible percent occurrence of mouse-like rodents, espe-
cially Brandt’s vole, in stomachs and feces, is probably explained
by the small numbers of voles present in the years when the ma-
terials were collected. Occurrences in summer feces of Mongolian
gazelle and domestic animals may be interpreted as the remains of
wolf meals and carrion. Observations exist of corsac foxes catch-
ing fish in the summer of 1944, in the dried-up lakes on the floodplain
of the Argun’ river (Brom, 1952). In half of all the examined
stomachs of corsac foxes and in 25% of their feces are found
insects, though as separate individuals. Of plant food, fruits of
sweetbrier were found.

In Trans-Baikaliya, the winter ration of the corsac fox hardly
differs from summer (see Table 36). Only in individual years, dur-
ing low rodent populations, feeding on carrion and various refuse
grows. In winter, a great quantity of tarbagan marmots are found
in stomachs and feces of corsac foxes (up to 51.1% occurrence).
Apparently, at this time, corsac foxes feed on carcasses of tarbagan
marmots, which die in summer burrows and were preserved there
by the cold (Cherkasov, 1884; Brom, 1952). Through tracks, it was
observed how the corsac foxes dig out from the snow the

295

430

entrances to abandoned marmot burrows and enter them. Frequent
feeding by corsac foxes on carcasses of marmots indicates that in
these regions, intensive epizootics occur among rodents. Occur-
rences of remains of Mongolian gazelle and roe deer are explained
in that the corsac fox in this region, as in Kazakhstan, feeds on the
remains of wolf prey and on carrion.

In the Mongolian People’s Republic, Brandt’s vole constitutes
the food base of the corsac fox in summer and winter, while other
rodents (gerbillines and hamsters) are of secondary importance.
Thus, on May 24, in the Choibalsan region in a burrow with a litter,
91 carcasses of mammals were found, of which 80 were Brandt’s
voles. In a burrow found on July 9 near Ulan-Tsirik, among 46
animal carcasses, 45 belonged to this same vole (Kucheruk and
Dunaeva, 1948). In Kentei aimak*, in two corsac fox burrows
excavated in May and June, remains of Mongolian gazelles, Cen-
tral-Asiatic [upland] buzzard [Вшео hemilasius] and horned lark
[Eremophila alpestris] were found, and in feces, the remains of
Brandt’s vole and birds. Brandt’s vole, within the limits of its range,
constitutes the main food of the corsac fox (Skalon, 1949). In the
stomachs of corsac foxes caught in spring and the beginning of
summer, Brandt’s vole, Daurian pika, quite a few orthopterous
insects and bones of small fish were present. The corsac fox eats
selitryanka fruits (Nitraria schoberi). More or fewer numbers of
remains of orthopterans may also be found in winter feces of the
corsac fox. Hunters believe that grasshoppers are a winter tidbit
for this carnivore; therefore, they already prepare them early in
autumn and use them in quantity as bait in traps. In southwestern
Khangai, in winter, corsac foxes also feed mainly on Brandt’s
voles, but also eat insects. Corsac foxes, especially young, feed on
orthopterans in other seasons also; with an abundance of locust, the
stomachs of the carnivores are literally stuffed with these insects
even during times of mass reproduction in Brandt’s voles (P.P.
Tarasov).

In the Mongolian Republic in winter, corsac foxes often feed on
carrion and refuse and closely approach populated places, appear-
ing even at the outskirts of Ulan-Bator [Ulaanbaator]. Sometimes,
corsac foxes become commensals with respect to diurnal raptorial
birds—upland buzzard (Buteo hemilasius) and saker falcon (Falco
cherrug milvipes), many of which winter in the northern part of

*Province—Sci. Ed.

296

431

the Mongolian Republic. Corsac foxes gather remains of their prey,
and at this time the stomachs of the animals become filled with hind
limbs, intestines and heads of voles; 1.е., all the parts which
buzzards and falcons leave rather than eat (P.P. Tarasov). Feeding
mainly on rodents, corsac foxes in the steppes and semideserts are
concentrated in places with an abundance of sagebrush voles or
Brandt’s voles, migrating there from the neighboring regions
(Nekipelov, 1935; Brom, 1952; Peshkov, 1954). They also gather in
regions where epizootics are spreading among rodents, or where
mass mortality has occurred among wild ungulates and cattle during
a prolonged period of continuous snow cover [dzhut].

With insufficiency of mouse-like and other rodents, or when
they are unavailable, corsac foxes suffer from hunger, markedly
lose weight and even die. In search of prey, they undertake migra-
tions of considerable distances (see later). Corsac foxes may remain
without food for 7-15 days and maintain their activity. During the
course of such a period, they sometimes do not leave the burrow,
particularly if a trap is set at its entrance.

Corsac foxes held in captivity in autumn consumed the follow-
ing amounts of food: males—from 120 to 480 g, averaging 218—347
g (various individuals); females—from 100 to 395 g, averaging 245—
259 g (two individuals; Brom, 1952). In Betpak-Dala, in the animals
caught in December in the early morning hours or at night, the
weight of the contents of a fully filled stomach was equal to 100—
130 g. Taking into consideration that the corsac fox hunts twice
daily—at dawn and at night—its natural daily ration in the wild is
approximately equal to 200—260 g, comparable to the contents in
captivity. In order to be sated, it is sufficient to it to eat one pika
(200 g) or about five voles.

In Betpak-Dala, in individual stomachs, the following foods were
found: female, on May 17—intermediate ground squirrel (adult),
darkling beetles; subadult male, on October 8—¢ray lark [Calandrella
pispoletta], mole crickets (7); female, on October 18—+ted-tailed
gerbil [Meriones erythrourus], eastern sand snake; female, on
November 27—-saiga meat and fat, social vole [Microtus socialis]
(2); female on December 21—-saiga meat, red-tailed gerbil, lizard
[Phrynocephalus sp.]. In Trans-Baikaliya, in individual stomachs
of corsac foxes caught in January were found: 1) Michnoi vole
[Microtus michnoi], Daurian hamster [Cricetulus barabensis],
Daurian partridge, 10 pieces of linen rags, 2) Daurian hedgehog

432

[Mesechinus dauuricus], Daurian partridge, a piece of shoe,
3) gray [brown] rat [Rattus norvegicus], a vole, a hamster, a
shrew, and grass leaves (Skalon, 1936).

With abundant prey, many carcasses may be brought into or
near the burrow by corsac foxes (see above). In the foothills of the
Zailiisk Alatau, carcasses of yellow ground squirrels (5. fulvus)
were found at the burrows, but rarely other food. On 29 May 1955,
at one burrow, 8 carcasses of yellow ground squirrels plus 8 heads
and 4 tails, a tail of a small five-toed jerboa [A. e/ater] and the
body of a lark, were found; at a second burrow, on 9 July 1955, 15
carcasses and 8 heads of ground squirrels were found (Kh.
Kydyrbaev). Such accumulations of carcasses are usually known
as “reserve’’, but actually are the remains of collected food. Some-
times, in periods of famine, corsac foxes eat the carcasses of their
fellows (Brom, 1952).

In the desert, the corsac fox is more often encountered near
wells and springs with water, but they are not attracted to them by
the water itself, but by the abundance of food. Even P.S. Pallas
(1811) noticed that the corsac fox rarely drinks and can remain for
a long time without it. “In captivity, and when fed meat, corsac
foxes can do well without water, and if it 1$ offered to them in the
cage, they are completely unattracted to it” (Nasimovich, 1951).

Home range. The dimensions of denning and hunting areas of
the corsac fox are little known. In the Pri-Caspian lowland, when
corsac fox was common, one litter occupied an area of 30—40 km’.
Burrows were unevenly distributed around the territory, occupying
the more elevated parts (Kolosov, 1935). In the steppes of Tselinograd
district, one burrow occupied an area of 0.8 km’, and 1.9-3.7 km?
for one litter (Chirkova, 1952). In the same district, near the
Kulanutmes river (semidesert), in 1951, when corsac foxes were
common, there were 27 burrows in an area of 10 km’, 7 of which
had litters. In the following year, after a sharp decrease in the
corsac fox population, in the same place in an area of 2.7 km’, only
one of 13 burrows found was inhabited; one corsac fox lived in it.
In the Kurgal’dzhin Lake region, in the winter of 1959/60, at the
end of February in an area of | km? 5 inhabited burrows of corsac
foxes were found, and 6 of these animals also were observed there
(B.D. Abaturov).

In southeastern Trans-Baikaliya, in the first half of August, the
young of one litter lived within a radius of 300 m of the den burrow

433

and spent the daytime in one or another tarbagan marmot burrows.
In this area, the corsac pups played, ran from one marmot mound
to another and entered the occupied and unoccupied tarbagan bur-
rows. In these same steppes, corsac foxes bringing prey to the
pups were observed at a distance of 6 km from their den burrow
(Brom et al., 1948). In the latter case, the hunting region of the
corsac fox family must have occupied about 113 km’. In winter, in
1948, in Trans-Baikaliya one corsac fox occupied 3 km? (Brom,
1952).

Burrows and shelters. The corsac fox is a relatively weak
animal, has many enemies, and therefore, in open landscapes, it
must hide in burrows and other shelters. The corsac fox uses bur-
rows also as protection from low or very high temperatures. The
corsac fox 15$ a poor burrower, and cannot dig in hard ground.
Therefore, it adapted to living in foreign burrows: of marmots—
bobak [М. bobak] and tarbagan [M. sibirica] and large ground
squirrels—yellow [5. ful/vus}—teddish [S. major] and red-cheeked
[5. erythrogenys]. It easily enters the openings into marmot bur-
rows, but has to widen those of ground squirrels in order to dwell
there. Rarely, it occupies abandoned burrows of red foxes and
badgers; only in soft sand or clay-alkaline ground does it dig itself
simple and usually shallow burrows. Thus, in the Barabinsk forest-
steppe, living places of this animal are simple in construction. They
have | to 4 entrances not more than | meter deep. In the center
of passages or at their end, the den chamber is built, which has no
bedding throughout the whole year. In the forest-steppes, the corsac
fox adopts for its own shelter the burrows of red-cheeked ground
squirrels, widening them, or inhabits old burrows of red foxes and
badgers. There, it digs itself only a shallow shelter with a short
passage, known by hunters as “dnevki” [day rest]. They use these
only in summer for resting and for hiding in case of danger. In warm
times of year, it usually rests by lying under a shrub, or in grass. In
winter a corsac fox, caught by a snowstorm in the steppe, far from
burrows, lies in a hollow dug into a snowdrift and allows itself to be
drifted over. It rarely hides in straw- or haystacks. During snow-
falls, it does not leave the burrow for 2—3 days (Sludskii, 1930).

In the steppes of Severo-Kazakhstansk district, the corsac fox
is adapted to the burrows of ground squirrels or marmots. The
burrow depth here is 40-150 cm, being explained by the ground
water level and deep winter snow cover. In the Sovetsk region, 1.5

434

km from Lake Chagla, in sagebrush-grass steppe, the den burrow
of the corsac fox has 2 exits, the general length of the passage is
4 m with side branches of .70 and 200 cm. Maximum depth of
hiding passages is 40 cm. Dimensions of den chamber are 30 x 40
cm; in it bedding is absent. On the surface between entrances, the
animals clear a small area, known by hunters as a “tochok’.
In the semidesert of central Kazakhstan near the Kulanutmes
river, the majority of 38 burrows found were situated on the sum-
mits of ridges, part on gentle slopes and a small number in the
bottoms of valleys. Entrances to burrows are often oriented toward
the north-northeast or northwest, and only in individual cases to-
ward the southeast and other directions. Entrance diameter varies
from 20 to 30 cm, with the average equal to 22 cm. Thirty burrows
(79%) had only one entrance, 2 had two entrances, 4 had three and
only 2 had 4. The majority of burrows are simple, and have only
one passage ending in a chamber. Complex burrows, with several
entrances and side-burrows, were rarely met with. Passage length
ranged from 140 to 450 cm, on average equaling 240 cm (of 12

298 burrows). Greatest depth of litter in passages varied from 55 to 115

297

Fig. 110. Corsac fox burrow under rock outcrop in semidesert at Kulanutme
river, central Kazakhstan. 27 July 1951. Photograph by E.I. Strautman.

299

435

cm, the average equaling 79 cm (of 11 burrows). The upper portion
of the burrow passages went through soft loamy soil, the chamber
was always located in very hard gypsum of a clay layer, which
could be cut only by an axe. All burrows were constructed in old
ground squirrel shelters. Chambers had dimensions of 40 x 30 cm.
In none of these was bedding found at the beginning of August.

Soil thrown out by the animals from the burrows does not form
a high mound of elongated form in front of it as is observed at red
fox burrows, but is scattered evenly around the entrance. Thanks
to this characteristic, to detect a corsac burrow it is more difficult
than that of a red fox. Around the den burrow occur many feces
of both adults, and especially, young. Moreover, some meters from
the entrance, in specially made pits are 2—4 “latrines.” Near and
to the side of many burrows there are 1—4 “tochok’, on which the
young pups played. In three cases, the living places were distrib-
uted in groups of 3 to 4 burrows each, and each one cf the shelters
was a distance of 40 to 500 m from the other; they were connected
to each other by trails. Such groups of burrows belong to one
family. In the Kurgal’dzhin Lake region, in the river valley, all
corsac fox burrows also were located in groups on elevations which
are not submerged by snow melt-water. In other sections there,
although single burrows were encountered, most areas were unin-
habited (B.D. Abaturov).

At 100 to 300 m from the den burrow, temporary burrows
(“dnevki”’) were sometimes met, with a passage length from 50 to
120 cm containing no chambers. At the beginning of August, in 12
burrows and 4 “dnevki’, 7 young animals were found; in one
burrow—2, while in the remaining—-singles. Usually, 70 to 80% of
the burrows are uninhabited.

In semidesert, the corsac fox utilizes burrows throughout the
whole winter. The entrance into the burrow is always cleared of
snow, which forms a small hillock. At the beginning of winter, often
there are several entrances cleared of snow, but by midwinter the
corsac fox cleans only one entrance of snow. A burrow dug in the
Kurgal’dzhin Lake region had, in the middle of winter, a length of
2.5 m and depth of 1.5 m. To a distance of 40 cm, the passage was
covered by hoar-frost, but beyond that thawed soil was found (B.D.
Abaturov).

In Betpak-Dala, corsac fox builds more complicated burrows
than in the steppe and semidesert, in relation to shallow snow cover

298

436

Fig. 111. Old den burrow of corsac fox with 23 exits. Edge of Karakum
desert near Tashkepri on the Murgab, Turkmeniya, 23 April 1962.
Photograph by A.A. Sludskii.

in winter, high air temperature in summer, and loose ground which
is easily excavated. Here, burrows are located on gentle slopes of
ridges or on level plateaus. Of 6 burrows, one had 3 enirances,
two—6, the remainder—8, 10 and 14 entrances. A family with 5
young lived in a burrow with three entrances on level ground among
peanut bushes. A second burrow was dug on a gentle south slope
covered by sagebrush and peanuts near a small solonchak 300 m
from a spring with water. It possessed 14 entrances, distributed
over an area 7 m in diameter. The passages led to a den chamber
located at a depth of 120 cm. At a distance of 70 m was located
another burrow with 6 entrances and an area strongly beaten down
by the playing of the young pups. The family of these animals used
two burrows simultaneously, in that in the burrow with 14 entrances
the parental pair lived, and in the burrow with six entrances—8
young.

A group of three burrows was located on a slightly undulating
clayey plain covered with sagebrush. The first possessed 18

437

entrances and was uninhabited. Forty meter from it was located a
second burrow with 6 entrances used only periodically. At 20 m
from the latter, a third, with 10 entrances, was inhabited. In it at a
depth of 125 cm was located a den chamber, in which was caught
an adult female and 3 young animals. Near the burrow there were
two cleared “tochok’. In the passages of the burrow were found
carcasses of five intermediate ground squirrels, badly decomposed
and with masses of fly maggots, remains of a small jerboa [A.
elater|, larks and desert wheatears [Oenanthe deserti]. At the
entrance to the burrow, little excrement was found, but the remains
of three intermediate ground squirrels, two eastern sand snakes and
some larks and desert wheatears were found. In a small side-
burrow, 12 scorpions were found. A complicated burrow with 6
entrances was found in a solonchak on the edge of the Taukum
sands.

Burrows of corsac fox in the northern Caucasus are also situ-
ated in groups (Dinnik, 1914). In this region, with sandy subsoil, it
is easy for the animals to dig burrows, and they are built very
widely. Each complex burrow with many deep-lying passages oc-
cupied an area having a diameter of perhaps about 40 to 60 m in
diameter. Sometimes, such burrows were located near each other.
These complex burrows received the name “corsac city” from
hunters. They believed that they were inhabited by many animals
that lived in a sort of colony. If such vast burrows actually existed,
they were inhabited, as a rule, by one family consisting of a pair of
adult animals and their litter.

On Barsa-Kel’mes Island in the Aral Sea, corsac foxes lived
in the burrows of yellow ground squirrels, suitably widened by
them. The length of their passages reached 4 m and were situated
at a depth of 1—1.5 m (Sludskii, 1939).

In the steppes of southeastern Trans-Baikaliya and the northern
part of the Mongolian Republic, the corsac fox lives almost exclu-
sively in the abandoned burrows of tarbagan marmots (Cherkasov,
1884; Nekipelov, 1935, 1952; Brom et al., 1948; Nasimovich, 1951;
P.P. Tarasov). The corsac fox does not widen the tarbagan’s bur-
rows. Corsac foxes often visit occupied burrows of tarbagans (Brom
et al., 1948).

In southeastern Trans-Baikaliya, in the course of six warm
months, the average caught out of each uninhabited [tarbagan]
burrow was: 0.6 corsac fox in 1947, 0.2 in 1948, 0.1 in 1949

300

438

(Nekipelov, 1952). In the Mongolian Republic, the corsac fox some-
times lives in burrows of the Daurian ground squirrel [Spermophilus
dauricus| after widening them. Five burrows found here had from
1 to 5 entrances. A denning burrow had one entrance and a den
chamber of 40 x 30 cm, without bedding (Skalon, 1949).

Therefore, in clayey steppes and semideserts, burrows of the
corsac fox are relatively simple and usually not deep. In snowless
deserts, especially in areas with soft ground, burrows occur that are
especially complex, and den chambers are situated deep. Den cham-
bers are also located deep in southeastern Trans-Baikaliya and the
Mongolian Republic, where winters are very severe and snow is
scant. The corsac fox does not strew grass in the den chamber.
Everywhere, corsac foxes build their burrows on plains and on
gentle slopes of ridges and hills. Burrows do not occur in ravines,
on steep slopes or among thickets.

Many symbionts use burrows of the corsac fox. In inhabited
burrows, parasites (fleas) and nidicoles (flies, beetles and others)*
are numerous. Scorpions were also found. The shelduck (Tadorna
tadorna) and ruddy shelduck (Tadorna ferruginea) nest in aban-
don burrows. Renard’s viper (Vipera ursini) and other reptiles
were also encountered there.

Daily activity and behavior. The corsac fox was previously
considered an exclusively nocturnal animal. However, recent ob-
servations have shown that it mainly leads a crepuscular mode of
life, and diurnal activity occurs, especially during the period the
young are being fed, when food is insufficient, and in times of
winter frosts.

Due to its relatively imperfect thermoregulation, the corsac fox
is obliged to use burrows more often than other foxes to reduce its
activity. At times of strong freezes and wind, it remains in the
burrow for longer than does the red fox. In the Barabinsk steppe,
during strong snowstorms, corsac foxes did not leave the burrows
for 2—3 days. Adapting to cold, the whole litter often lives in the
burrow in winter. Because of this, 2 to 5 animals are successfully
dug out simultaneously from one burrow (Sludskii, 1930). Even in
severe freezes, the corsac foxes, being hungry must remain long
outside the burrow searching for prey. On calm frosty days, the
animal may be observed lying on the snow near the entrance to a
burrow, while the snow beneath it does not thaw.

*Nest inhabitants—Sci. Ed.

Fig. 112. Corsac fox burrow near Imam-Baba. Edge of Karakum, Murgab.
22 April 1962. Photograph by A.A. Sludskii.

302

440

The corsac fox inhabits semideserts and deserts, where air
temperature in summer rises to 40°C and even to 50°C, and the soil
surface becomes heated to 75°C and more. The corsac fox es-
capes from such high temperatures in burrows and in the warm
season, it chiefly leads a crepuscular mode of life.

The corsac fox is a carnivore, capable of capturing food only
in those places where there are no dense thickets (in summer) or
deep snow cover (in winter). The corsac fox is not adapted to
movement on snow. In the animals from Betpak-Dala, the length of
the forelimb from the chest to the ground ranges, in males, from 17
to 23 cm (average—19.0 cm; 6 specimens) and, in females, from 16
to 19 cm (average—17.3 cm; 9 specimens), while in the Karagan
red foxes obtained from the same region, corresponding values are
26.3 and 24.0 cm, respectively. In spite of low weight (2.0—2.5 kg),
the corsac fox has a great weight loading on the supporting surface
of the foot—61 g/cm? for male animals from Trans-Baikaliya
(Nasimovich, 1951). In animals from Betpak-Dala (males), it is
from 68.3 to 80.0, or an average of 75.2 g/cm? (4 specimens), and
in females it is from 43.6 to 74.0—an average of 54.4 g/cm’ (9
specimens). In Karagan red fox, this weight loading is 62 g/cm? in
males and 60 g/cm’ in females, and in red foxes from heavy snow
regions, it is less: in Lapland red fox—27—30 g/cm? (Nasimovich,
1948) and in Pechora red fox—28 g/cm? (Teplova and Teplov,
1947). Thanks to the great weight loading and relatively short limbs,
the corsac fox deeply sinks into loose snow, and with snow cover
depth of 15—20 cm, it moves with great difficulty. At that time, the
corsac fox stays in burrows or uses trails and tracks of other
animals and humans. In distinction from the red fox, the corsac fox
does not have a “mouse-leap’—a unique jump on rodents found
under the layer of loose snow. Therefore, with deep snow cover,
corsac foxes seek out areas where the snow is blown off or com-
pacted by wind, and they also are restricted to places of winter
grazing by large herds of saigas and domestic livestock. In obtain-
ing food from under the snow while winter grazing, these ungulates
dig up and trampling it, help the corsac fox to move and to catch
rodents (Skalon, 1952; A.A. Sludskii). In the presence of deep
snow cover, hard-crusted or ice-covered ground, the animals suffer
from hunger and leave that region.

In places with tall grass cover (meadow grass) as, for instance,
in Badkhyz (southern Turkmeniya), corsac foxes constantly use the

301

441

numerous trails beaten by goitered gazelle and Кап (V.G. Heptner).
In winter, corsac foxes were observed at carrion during the day,
and on windless sunny days, lying on the snow near a burrow
(Sludskii, 1930; B.D. Abaturov). In Betpak-Dala, corsac foxes were
met with in December around 10—11 in the morning, and during the
afternoon, when they approached domiciles to feed on refuse. At
that time, they were also active after dark. The same thing is
known for southeastern Trans-Baikaliya and Mongolian People’s
Republic (Brom et al., 1948; Skalon, 1949, N.V. Nekipelov). In the
northern part of the Mongolian Republic, the corsac fox in winter
begins hunting at the end of the day, before evening and scurries
about searching for food the entire first half of the night; it again
hunts early in the morning, and very rarely during daytime.

In summer, corsac foxes are also active not only at twilight and
at night, but often hunt during the day, especially when they are
feeding young (Nasimovich, 1951). In captivity, at air temperature
from 3° to 10°C, corsac foxes were active at 1900, 2300 and 0200
hours [Fig. 113]. With reduction of air temperature from 25 to 0°C,
activity of the corsac fox decreased threefold (Kalabukhov and
Poluzadova, 1946; Kalabukhov, 1950).

Young animals, 1.5—2 months of age, assume a more diurnal
mode of life. Thus, in Betpak-Dala, from 11 to 27 May, a litter of
8 pups might be observed on the surface already at 5 in the morn-
ing. Until 9 am, the corsac pups played near the burrow entrance,

SUAS NIM NG 22 ТЗ 29
Hour of the day

Fig. 113. Daily cycle of activity of corsac fox in an experiment at 3—10°С
(after Kalabukhov, 1950).

303

442

moving not more than 5 meters away. When it became hot, they
hid in the burrow and by 11 am only individual animals remained on
the surface, but in the course of the whole day 1 to 2 corsac pups
usually were lying near the burrow, sometimes changing places. At
the end of the day, the corsac pups came out from the burrows
more frequently and remained on the surface for a longer time. By
20:00 hours, the whole litter was already inside the burrow. Coming
out of the burrow, the corsac pups looked around for a long time,
moved slowly and often scratched; apparently, they were greatly
disturbed by fleas. At sunset, they cautiously began to play. When
threatened by danger, they broke off playing and stretching their
necks, endeavored to see what had frightened them. When afraid,
the pups barked, making a sound which may be given as “vyau-
vyau”. After quieting down, the corsac pups again returned to play
which continued until the approach of full darkness. But even later,
at 11, 12 and 1 at night, when the burrow was illuminated, the
whole litter could be seen near it.

When coming out of burrows, the corsac fox at first thrusts out
its head only, then shows itself to the shoulder, and only after being
sure that it is quiet all around, does it come out on the surface.
After coming out, the animal stretches itself several times, looks
around, and then sits down, or sometimes lies down, near the bur-
row, continuing to observe its surroundings. Sensing danger, it sits
and watches closely, slightly nodding its head. If the alarm is shown
to be not false, it begins “barking” in that direction, continuing,
however, to sit in its place. After quieting down, with the approach
of twilight, it sets out on the hunt and already at 21:00 hours, the
animal could be observed beating back and forth along the road for
jerboas.

Both in summer and in winter the corsac fox, lying near the
burrow entrance, does not respond.to a human approaching it be-
fore a distance of 30 to 50 meters; after that, it runs aside and does
not run into the burrow. When followed, it runs to a neighboring
burrow, and remains motionless. Thus, even injured animais behave
in the same manner, but when escaping from a wolf or eagle, the
corsac fox enters the first available burrow, even if inhabited by
marmots.

It is interesting that the corsac fox has the ability to remain
motionless, feigning death. The animal is sometimes found thus in
an excavated burrow. It lies completely without movement, relaxed

443

and with its eyes closed. Taken up in the hands, it remains in this
state, completely lifeless. But, if it is put on the ground, it opens its
eyes, jumps up and runs away. (Similar behavior is known for red
foxes, jackals, striped hyaenas, ratels, raccoon dogs and especially
opossums.)

The corsac fox is somewhat timid and cautious. It permits
humans to approach as near as 10 meters, and then runs off for
100—200 т and remains motionless. It also permits a car as close
as 5—6 meters. It is caught in traps more easily than the red fox.
With abundant food, for example, at a time of massive death of
ungulates from starvation, it sometimes remains with red foxes and
even dogs near carrion at cattle burial grounds and dumps (Sludskii,
1930).

In summer, corsac foxes hunt alone: the animal slowly walks
against the wind and on smelling or seeing prey, conceals itself by
pressing to the ground and running from one cover to another. Not
far from the intended victim, it jumps up and overtakes it in a few
springs. Rarely, it runs at a slow gait, and startling its prey, for
example, sagebrush vole, pursues its prey, and overtaking it, en-
deavors to pin it with its front feet. Where jerboas are numerous,
it watches for them by remaining motionless behind bushes along
roads or paths, where these rodents run, at the approach of twilight
or at sunrise. On the ground, it searches for nests and preying on
nesting birds, eats the nestlings and eggs. It catches goslings when
their parents lead them to the steppe to feed, and ducklings at the
time the adults lead them from the nest to water or transfer them
from one lake to another. The corsac fox hunts lizards by jumping
on little shrubs under which they are hidden, and endeavoring to
roust them into the open.

In winter, it also searches for food singly. In the literature,
there is some information stating that corsac foxes sometimes hunt
in pairs or even in small packs. Actually, they gather as several
individuals around large carrion, but approach it singly, following the
tracks or trail trampled by other animals.

In the search for scarce winter prey, the animal goes at a trot,
stopping and sniffing. Scenting or hearing the squeak of an animal
under the snow, it quickly digs the snow trying to catch the rodent.
With snow cover deeper than 10—15 cm, or when it is compact or
covered with a hard crust, the hunt of the corsac fox is usually
unsuccessful.

304

444

Согзас fox possesses а good sense of smell. One can success-
fully follow by tracks on the snow how it turned at a right angle to
its path and 20—30 m to the side dug out from under the snow ап
old sheep leg, horse hoof or leather scrap (Brom, 1952). The corsac
fox preys on willow ptarmigan and gray partridge night-roosting in
the snow, black larks and snow buntings, and watch for white hares
on their trails. In breaks in the reeds, the soil buried under the snow
is not frozen, and it catches water voles by excavating their burrows.

Corsac fox tames well, and can live in captivity for a long time.

Seasonal migrations and transgression. The approach of
cold, the entrance into hibernation of rodents, the torpor of reptiles
and insects, the migrations of a series of birds and the falling of
snow force the corsac fox, in late autumn or the beginning of
winter, to migrate to the south, but sometimes also in other direc-
tions. Obtaining winter food, which is rare itself, becomes more
difficult if snow cover is deep—more than 15 cm, or forms а hard
snow crust or ice-covered ground. Abundance and availability of
food depend also on other factors: severe frosts at the beginning of
winter when there is still no snow, steppe fires, migration of ungu-
lates, etc. All these phenomena often take place in the forest-steppe,
steppe, and some in the northern deserts of Kazakhstan.

In individual winters, the number of corsac foxes in the steppes
increases from the tally of animals arriving from the forest-steppe,
and with low numbers or difficulties in capturing rodents in the
steppe, corsac foxes leave it for the semidesert. In the northern and
central parts of its range, including the northern deserts, the sea-
sonal migrations of these animals bear a more or less regular
character and take place almost annually. In the beginning of win-
ter, the majority of corsac foxes abandon summer places and search
for sections richer in food (Polferov, 1896). In the steppes of north-
ern Kazakhstan, in the heavy snow winter of 1940/41, corsac foxes
(and also wolves and red foxes), migrated en masse to the south-
west to the Turgai river and were encountered in Priishmisk and
other regions of Severno-Kazakhstansk district, where they had not
been previously observed for many years (I.R. Zenkin). In the
winter of 1942/43, in these same places, corsac foxes migrated to
the north. One hunter there caught 56 animals in the season, at a
time when, in the usual years, only 2—6 animals were caught (Е.Е.
Chernov). In 1947 (in November), intensive emigration to the south
of corsac foxes was observed there, connected with the low

445.

number of mouse-like rodents in the places of their summer resi-
dence. In the heavy snow winter of 1948/49, massive emigration of
corsac foxes was recorded in Kustanai district. The animals went
to the south, while from the steppes of Tselinograd district, some
migrated to the south and others to the northeast, towards Om’sk
and Pavlodar. As a result, corsac fox population decreased by
about 40% (K.T. Popov). In 1950/51, in connection with the early
formation of snow cover, which was very compact as a result of
thawing and refreezing, forming a thick frozen ice-crust, from the
middle of December massive emigration to the south was observed
from Kustanai, Tselinograd and Karaganda districts.

Such migrations of corsac foxes, having in different years vari-
ous directions and intensity, depending upon weather and food
abundance, were also observed in other parts of the range. By the
middle of December, with the formation of a deep snow cover in
the semidesert, the population of corsac foxes in Betpak-Dala in-
creases as a result of migration from the north. At that time, their
tracks are encountered in great numbers and the animals them-
selves may often be observed. In the northern and central parts of
the Betpak-Dala desert, with the formation of the snow cover, the
corsac foxes begin to suffer from a great insufficiency of food,
since of the rodents, only the red-tailed gerbil remains active and
they rarely come out of their burrows at that time. Therefore,
corsac foxes here in winter migrate after saiga herds, feeding on
the carcasses of antelope killed by wolves or starvation. The ani-
mals move at night. Each year in December, saigas move to the
southeast for 4 to 5 days through the central region of the Betpak-
Dala; in this short period or a bit later, the corsac foxes also
migrate. By the end of December, the majority of corsac foxes
have already followed the trail of the saigas southward to the Chu
river valley. Those remaining suffer greatly from hunger. There are
no rodents, and, at this time, the remains of saiga carcasses killed
by wolves, foxes and corsac foxes also disappear. The last animals
remain without prey and are also forced to migrate, following the
saigas. Moreover, in places of massive saiga concentrations, the
snow is much beaten down and trampled by them, which facilitates
the movement of corsac foxes and their capture of rodents. Many
corsac foxes die from starvation and wolves. In winter in Trans-
Baikaliya and the Mongolian Republic, corsac foxes follow ‘the
migrating herds of Mongolian gazelles (Brom, 1952 and others).

305

446

The cause of corsac fox migrations may also be steppe fires,
which cover great areas. Migrations explain their appearance in the
regions, where in summer they are absent or rarely met with. For
instance, in a series of regions in Kokchetav district, they are found
only in winter, appearing there at the end of December. The mag-
nitude, and often also the distance of migration, depend upon the
number of corsac foxes and the degree of change in conditions
which induces migration (much snow, frozen crust, cold, steppe
fires, etc.).

Corsac foxes migrate also in the east—in Pri-Baikal, Trans-
Baikaliya and the Mongolian People’s Republic. Thus, in some
years, they appear in the Selenga river valley in Kyakhtinsk district,
penetrating there from the MPR (Fetisov, 1949; Surmach, 1957). In
southeastern Trans-Baikaliya, during the time of winter migration,
corsac fox sometimes transgress very far to the north and appear
in forest-steppe regions (Aleksandrozavodsk region—Algachi,
Man’kovo, Bokhto and Bazanov; Stretensk region—Kuenga river,
left tributary of the Shilka; E. Pavlov, 1949; N.V. Nekipelov). In
the last decades, mass appearance of corsac foxes in Trans-Baikaliya
was noticed in 1946—1947 and in 1955. The main reason for the
mass appearance of corsac foxes in Trans-Baikaliya was their
migration from the northeastern part of the Mongolian Republic,
where in 1942/1945 and in 1953 there was in places mass repro-
duction of Brandt’s vole, which later disappeared, that provoked the
mass migration of corsac foxes northward (N.V. Nekipelov). In
Trans-Baikaliya, corsac foxes migrate, moving out from regions
where Brandt’s vole and Daurian pika populations are reduced.
In Trans-Baikaliya, the corsac fox migration is more rarely caused
by deep snow cover (Cherkasov, 1884; Dorogostaiskii, 1935,
Podarevskii, 1936).

Extinction of rodents or other unfavorable conditions cause mass
migration and settlement of corsac fox even beyond the range
boundary. In this way, they sometimes populate new territories, but,
as a rule, not for long. Thus, in separate years corsac foxes ap-
peared in the middle Volga region, where they were hunted in 1930
and 1933 in the Bol’sheglushitsk region. They also transgressed in
1936 into the Bol’shechernigovsk region (Dobrokhotov, 1937).

In the severe heavy-snow winter of 1955/56, corsac foxes
appeared in the forest-steppe of the Volga region at Vol’sk city and
Teplovka village, arriving, apparently, from the Trans-Volga steppes.

306

447

Transgressions hither of corsac foxes there are so rare that none
of the old-timers remembers such an immigration (Kirikov, 1959).
In the winter of 1943/44, corsac foxes were encountered in
Oktyabr’sk, Bugul’minsk and Bavlinsk regions of the Tatar ASSR,
where they had not previously been observed (Popov and Lukin,
1949). In order to reach the regions mentioned, the animals must
have traversed not less than 320 km in a straight line. In the 30’s
of this century corsac foxes were caught in the Beloretsk region in
Bashkiria, 280 km from the border of their permanent range. In the
past century, corsac foxes appeared beyond the Ural range, in the
Shadrinsk region, 240 km north of places where they usually occur
(Sabaneev, 1874). In the winter of 1948/49, when the snowfall was
heavy in Kazakhstan, corsac foxes were caught in Sorochinsk,
Perevolotsk, Krasnokholmsk, Orenburgsk, Burtinsk, Troitsk and other
regions of Orenburg district, 200 km and more to the north of the
core habitat (Darkshevich, 1950). In 1952, there were many corsac
foxes in Chelyabinsk district, whence they arrived from Kustanai
and Severo-Kazakhstan districts. The same was also observed in
western Siberia, where the corsac fox penetrated northward to
Lebyazh’ir in Kurgan district, Tyukalinsk, Tara and Vengerov which
are 50-200 km beyond the usual northern limits of their distribution
(Shukhov, 1928; Kirikov, 1959, 1960).

Massive emigration of corsac foxes from the steppes of
Kazakhstan to the forest-steppe of western Siberia took place at
the-end of the 10’s and start of the 20’s and especially in the 40’s
of the current century. The First World War and the Civil War
which followed it distracted many hunters from the fur harvest.
The corsac fox catch was greatly reduced. At the same time, the
fields were badly treated. It was not possible to harvest and thresh
the ripened wheat; much arable land was abandoned, allowing mass
reproduction of rodents in them. Under-harvest of corsac fox and
the wealth of food reserves favored its reproduction and intensive
colonization beyond the usual limits of the range. In 1917, corsac
foxes appeared in great numbers in the Barabinsk steppe east of
the Irtysh. Their number increased without pause until 1921. There-
after, their population began to decrease, and by 1929 they
completely disappeared (Sludskii, 1930).

In the Altai territory, in the territory of the Kiprinsk region,
corsac foxes were common up to 1916, then disappeared. In the
winter of 1920/21 in this region, a mass appearance of this animal

448

was noted, moving to the north. By the end of the 20’s, corsac
foxes disappeared there (Velizhanin, 1931). The same picture was
also observed southward—in Semipalatinsk district (Selevin, 1930).

Migration of corsac foxes also takes place in deserts, where it
is associated with prolonged, continuous snow cover accompanied
by mass death of ungulates. After the heavy-snow winter of 1949/
50, corsac foxes appeared in great numbers south of Zapadno-
Kazakhstansk and Aktyubinsk districts and penetrated to the
northern shore of the Caspian. Already in December 1948, many
of these animals arrived from the north in the Syr-Dar’ya river
valley (Sludskii, 1953). The corsac fox passed through from the
north in significant numbers to Dzhambulsk and Alma-Atinsk dis-
tricts and to the Chu valley. They appeared not only in the foothills
but also low in the mountains. With the formation of deep snow
cover, they began to return again to the sands of the Pri-Chuisk
Muyunkum, but there was also much snow there, and the animals
died of starvation.

At the time of emigration from the northern deserts, corsac
foxes crossed the ice of [Lake] Balkhash and appeared in southern
Pri-Balkhash (winters of 1938/39; 1943/44, 1945/46, 1948/49 and
especially 1950/51). In the foothills of the Dzhungarsk Alatau and
in the Alakul’ depression, corsac foxes appeared in 1938/39, 1947/
48 and 1948/49, i.e. in nearly the same winters as in southern Pri-
Balkhash. In Zaisan depression, the corsac fox population sometimes
increased on account of the animals emigrating hither from Xinjiang,
which took place in November—December 1938. During the time of
emigration of corsac foxes from northern-type deserts southward
from the southern edge of the semideserts to the foothills of
the Tien Shan, they traveled a straight-line distance of 400 to
600 km.

With food scarcity or difficulty in obtaining it, corsac foxes
approach populated places, even large cities. Thus, in the severe
winter of 1939/40, in Pavlodarsk district, they were encountered in
great numbers near settlements, and regularly fed on meat-packing
wastes near the city of Pavlodar. In the heavy-snow winter of
1940/41, in Severo-Khazakhstansk district (Sovetsk region, near
Barankul’ settlement), corsac foxes appeared in such great num-
bers that one hunter obtained more than 50 animals in the season.
In the winter of 1948/49, in Karaganda district, corsac foxes and
red foxes often appeared on the outskirts of settlements and within

307

449

the precincts of Karaganda, where they fed on rubbish; several
emaciated animals were caught by hand.

Therefore, for the corsac fox in its whole range, it is charac-
terized by considerable displacement, governed, first of all, by periodic
food deficiency. In the forest-steppes, steppes, semideserts and
deserts, the corsac fox performs migrations almost yearly. The
direction of migration changes, but more often the animals go south-
ward, rarely to the north or other directions. Distances traversed by
corsac foxes during times of migration are not established, but the
boundary of the range, in connection with emigration, “pulsates”
within limits of 50-600 km. As a result of migration and emigration,
the corsac fox population in one or another region may change as
much as ten fold.

Reproduction. The corsac fox is monogamous, the male also
participating in nurturing the young. Apparently, having formed a
pair, corsac foxes live together until one of the members dies. In
the Moscow zoo, a male living with two females in one cage
covered [mated with] only one of them. In the course of a year
only one litter occurs.

In Kazakhstan, in the steppe zone, rut in these carnivores starts
in February; in the semidesert and desert zones—in January or the
beginning of February; and in the steppes of southeastern Trans-
Baikaliya, in February. At the time of rut, in the tracks orange-red
spots are often met with, left in those places where the animal
urinated. Two corsac foxes usually live near some burrows. The
animals usually rut at night; at this time their peculiar barking is
often heard. Several males run behind a female in heat for the first
time, or for one left barren for some reason, and fight usually
occurs between them. The female usually mates in the burrow.

The duration of pregnancy is not accurately established, prob-
ably equaling 52 days on the average. The earliest litter was found
in Aktyubinsk district on 15 March 1953. In central Kazakhstan, at
Lake Tengis, on 11 April 1957, a female was caught with 12
embryos, the body length of which equaled 10 cm and the average
weight, 30 g. In Kustanai district, the first litter was found on 25
April, and between 5—15 May the young corsacs came out from the
burrow. In Kokchetav, Tselinograd and Ural’sk districts, the young
appeared at the burrows оп 15—25 April. In Semipalatinsk and
Vostochno-Kazakhstansk districts, they begin to emerge on 5—10
April. In southeastern Trans-Baikaliya, a recently parturient female

450

was caught on 16 April 1961 (N.V. Nekipelov). Litters here appear
at the end of April-beginning of May (Peshkov, 1954). In the
Moscow zoo, pupping of the corsac fox was observed eight
times—between 18 March and 10 May. From Table 37, one may
judge litter size, and when young began to emerge from the burrows.
On 10 June 1942, in the Zharminsk region of Semipalatinsk district,
16 young of the same age were dug out from a burrow.
Therefore, in Kazakhstan, there may occur in a corsac fox
litter from 2 to 16 young, but more often 3—6 (average of 5 for 100
samples). In the northeastern Cis-Caucasus, three young animals
were found in a burrow (Heptner and Formozov, 1941). In the
ephemeral deserts of Uzbekistan, the litter usually consists of from

Table 37. The number of young in one litter of corsac
fox in Kazakhstan

Total Average
of pups in litters number

litter in litter
Zones

and districts

True steppes:
Kustanaisk, ово] 29 4.9
Зеуего-
Kazakhstansk,
Pavlodarsk
[districts]

Dry steppes:
Ural’sk — 23 5 2 —— — — 12 4.6
Aktyubinsk

Kokchetav and — 8 8 6 10 3 2 1 — 38 5.0
Tselinograd

Semideserts:
Karaganda — —— — 41°55 91 Se 11 6.3

Vostochno- — 2 1 4 — 2 — — — 9 4.9
Kazakhstansk

and

Semipalatinsk

Deserts:
Dzhambul ом SS м о 1 3.0
Total Le 2S le 2012105 1 1 100 5.0

in %. Я ee pele] 100

308

451

3—4 young, but many of them die while still in the burrow. The
reason for juvenile mortality in the burrow period of life is believed
by many hunters to be due to ants which attack the still-helpless
animals. They also point to the possibility of diseases (Suleimanov,
1939). In the steppes of southeastern Trans-Baikaliya, the litter
contains from 5—7, and up to 10, animals (Cherkasov, 1884; Peshkov,
1954). In the Mongolian Republic, in the Kerulen valley on 22 May
1944, 7 young were found in an excavated corsac fox burrow
(Skalon, 1949), and in Choibalsan region, on 24 May—11 pups half
the size of adults (Kucheruk and Dunaeva, 1948). In the Mongolian
Republic, the number of young animals in the litter varied from 2
to 12 (Bannikov, 1954). In the Moscow zoo, in five litters there
were 3, 4, 5, 6 and 7 young.

It is not known whether fertility of corsac foxes increases
significantly in years of food abundance as happens, for example,
in arctic foxes, although the larger litters were found in the years
with abundant sagebrush voles or Brandt’s voles.

Growth, development, and molt. Corsac pups are born blind
and with closed ear passages. They are covered with a light brown
fluffy pelage. The tail is monocolored, while in the red fox its tip
is always white. The tone of their fur becomes increasingly more
and more yellow with age.

Weight of newborns is 60—65 g; body length, 13.5—14.0 cm.
They can see on day 14—16. At 28 days of age, they begin to eat
meat (E.A. Petrova). They grow rapidly. Thus, in central Kazakhstan
young animals—females caught on 28 May 1960, weighed 245—262 g
and of corsac pups from a litter captured on 29 May 1960, males
were 392—400 g and females, 378-405 g. In this same region, two
young females caught on 31 July 1951 each weighed 1900 g al-
ready, and a male caught on 3 August 1951—2050 g. In the
Betpak-Dala desert, a young female caught on 11 May 1955 weighed
1250 g, and a male caught on 17 May 1956—1200 2. Young females
also captured there weighed: 20 September 1957—1800 g, and 13
December 1957—2000 g. In Turkmeniya, on the southern edge of
the Karakum, a young animal caught on 6 May was half the size
of an adult. Weight of such an adult female is 2.3—2.5 kg, and of
male, up to 2.6 kg. Consequently, at the beginning of May in
Turkmeniya and the middle of this month in the northern deserts,
young animals already have attained half the dimensions of adults.
Farther north, they are smaller in May. However, already by the

309

452

beginning of August, the young weigh 1900—2000 g, 1.е. slightly less
than adults, even in the semidesert. Apparently, the corsac pup
completes its growth in 4—5 months.

In the den burrow, the female lives excellently with the pups
until they are 2 months of age. The male, though he participates in
the raising of the young, apparently rests in another burrow or
outside it, since in no case has he ever been found in any shelter
with the pups (sample size is still small). When the pups approach
full growth, the female sometimes lives with the male in one bur-
row, while the young are in a neighboring one.

In spring and summer, many fleas are found in occupied corsac
fox burrows (Brom et al., 1948), which greatly bother the animals.
Apparently, because of the parasites and dirt in the burrow, the
female changes burrows 2—3 times, carrying the young with her.

The young disperse early. Out of nine burrows with young,
excavated in June—August, in only one burrow were two young
revealed, while in the remainder, only one. Young and adult corsac
foxes also change burrows often.

In burrows excavated in November—December, 2—5 young ani-
mals were often found (Dinnik, 1914; Sludskii, 1930, 1939; Heptner,
1932). On Barsa-Kel’mes island, in December, three young males
were caught in one burrow, and in a second—three males and one
female (Sludskii, 1939). Apparently, with the onset of cold, the
earlier dispersed young corsac foxes gather again in one burrow.
It is not known whether they live in groups.during the course of the
whole winter, though there are references to litters of corsac foxes
that separate only at the time of onset of rut (usually in February)
or in spring. Assertions that several adult animals may live peace-
fully in one burrow require verification.

Duration of lactation is not established; a female caught in
Betpak-Dala on 17 May in a burrow with three young that weighed
1200 g already had no milk in her glands and her teats were dry.

Females attain sexual maturity at the age of 9-10 months. In 28
animals investigated at the age of 4 months, there were 53.4%
males and 46.6% females. Under favorable conditions, the corsac
fox becomes more fertile and therefore its numbers may quickly
increase.

It is accepted that the corsac fox molts two times per year—
in spring and autumn. In spring, the winter pelage is completely
replaced; it 15 not established whether the pelage is also completely

453.

replaced in autumn or not. By the end of winter, the hairs lose their
luster; gradually dulling, the guard hairs become dry and frayed and
their ends bend. First of all, fur on the head and limbs 15 replaced,
then fur on the nape and shoulders molts, later the anterior part of
the back, side and thigh, then the posterior part of the back and,
last, the rump and tail. Consequently, in spring the molt proceeds
from head to tail. Spring molt lasts quite a long time. In the northern
deserts (Barsa-Kel’mes Island), it is already initiated by February
15 and lasts until the middle of May. In a nursing female caught on
17 May 1956 in Betpak-Dala, the winter fur was nearly fully shed,
and was retained only on the sides, rump and tail. Animals caught
in this desert in August had summer fur, while those captured in the
beginning of November well exhibited new fur, but it was still not
mature. The growth of the [winter] pelage ends only at the end of
November, just after severe cold makes its appearance.

Consequently, growth of winter fur is in reverse order. First
growth commences on the tail and rump, then on the thighs and
back, further on the nape and sides, and lastly on the paws, belly
and head.

Enemies, diseases, parasites, mortality and competitors.
Everywhere within the boundaries of the Soviet Union and the
Mongolian Republic the competitors of corsac fox as regards food
are the wolf, red fox, and steppe polecat [Mustela eversmanni]
and in the forest-steppe and steppe zones, also ermine and weasel;
in the Barabinsk steppe and in Trans-Baikaliya—kolonok [М.
sibirica] and in the latter region the solongoi [M. altaica] as well. In
central Kazakhstan and in Trans-Baikaliya among the competitors of
the corsac fox is also considered the manul [Felis manul] and
in the deserts—marbled polecat [Vormela peregusna], steppe [F. 5.
libyca] and sand [F. margarita] cats, jackal and several others.

In the spring—summer period, wolf feeds mainly on the same
rodent species on which the corsac fox feeds, and is very common
in some places. By comparing fur tanning of both species, one may
see that in the Tselinograd district, for each corsac fox caught,
there was 0.5 wolf and in Karganda district, 1.4 wolves. Red fox
is an especially dangerous competitor for corsac fox. Throughout
the whole year, it feeds on the same food as corsac fox. It is
stronger than the corsac fox and more successfully catches rodents
of moderate and large size; it is also better adapted to mousing in
regions where the snow cover reaches 10 cm and more. Only in

310

454

the semideserts does the corsac fox, apparently, hunts more suc-
cessfully than the red fox. The red fox is ecologically more plastic
than the corsac. In northern Kazakhstan, judging by fur tanning
data during recent years, the red fox is numerically predominant
over the corsac fox. Thus, in Kustanai district, 2000—3000 red foxes
are caught and only 500-800 corsac foxes, in Severo-Kazakhstansk,
2000—2400 red foxes and only 100—300 corsac foxes, and in
Pavlodarsk, up to 2500 red foxes and 1000—1500 corsac foxes.

In semideserts, before ploughing of virgin soil, corsac fox ob-
viously predominated over red fox. In Tselinograd district, for
example, in the 30’s of the current century, only some tens of red
foxes and more than a thousand corsac foxes were caught. In the
40’s, already 400—500 red foxes were caught and 1000—1200 corsaa.
In the 50’s, the population of red foxes greatly increased and to-
gether with this the maximum catch; in a season, 2900 fox skins
were prepared and at the same time 2400 corsac fox skins. There-
fore, competition between these two species of carnivores rapidly
increased. In northeastern Cis-Caucasus, in the season of 1924/25,
one hunter took 15—30 corsac foxes and 10—20 red foxes (Heptner
and Formozov, 1941).

In the northern deserts, the red fox population was always
much higher than the number of corsac foxes. According to tanning
data, in Kzyl-Ordinsk district, 4 to 5 thousand red foxes are caught,
maximum 8 thousand, and corsac foxes—200—400, up to 900 indi-
viduals. In Alma-Atinsk district, the corsac foxes are still fewer:
during the last ten years here, the maximum catch of red foxes
reached 15 thousand, and corsac only 650 individuals. In the steppes
of Trans-Baikaliya, corsac fox populations are predominant over
red fox.

Therefore, in virgin dry steppes and semideserts, corsac foxes
predominate, and in deserts—red foxes. Ploughing of virgin lands
leads to disappearance of corsac fox and growth in numbers of red
fox, of which the Ukrainian steppes, and in recent years—also
Kazakhstan, are obvious examples. Competition between red and
corsac foxes for food becomes particularly intense in autumn and
winter, when the majority of rodents enter hibernation, many birds
migrate, and reptiles and insects are torpid.

Throughout the range of the corsac fox, the steppe polecat is
numerous, also a serious competitor of the corsac fox. In the 30’s
of the present century in Kazakhstan, up to 500 thousand skins of

455

this carnivore were taken every year; 1.e. one hundred times more
than corsac foxes. The intensity of competitive relationship be-
tween these animals can be partially judged by the frequency of
their capture in traps at tarbagan marmot burrows. For a series of
years in eastern Trans-Baikaliya, during the warm months, 105
corsac foxes, 4 wolves, 9 weasels, 67 solongoi, 412 steppe polecats
and 8 badgers were caught in traps set for marmots (Brom et al.,
1948). For every corsac fox, four polecats were obtained. Ermine,
weasel, kolonok, solongoi, marbled polecat and wild cats, because
of their relatively low number and small size, are not considered as
serious competitors of the corsac fox. The corsac fox itself rarely
attacks small carnivores.

Among avian competitors of the corsac fox are manifested
saker falcon (Falco cherrug), pallid and northern harrier (Circus
macrourus and C. cyanus), steppe eagle (Aquila rapax) and
rough-legged buzzard (Buteo lagopus). In the western range, the
long-legged buzzard (Buteo rufinus) appears as a competitor of
the corsac while in the east—in Trans-Baikaliya and the Mongolian
Republic, it is the upland buzzard (Buteo hemilasius). The steppe
eagle and, rarely, the upland buzzard are enemies of the corsac fox
and with regard to saker falcon and common buzzard [Buteo buteo],
this animal becomes a commensal (see above).

The wolf is the greatest danger to the corsac fox. In the sec-
ond half of winter, especially with much snow, wolves kill many
corsac foxes (Velizhanin, 1931). Thus, in Tselinograd district, in the
winter of 1947/48, in February, one hunter found six corsac foxes
torn apart by wolves. In Semipalatinsk district, in March—April 1948/
49, a hunter found the remains of 10 corsac foxes eaten by wolves,
and in the winter 1950/51, in the Kurchumsk region, in one day,
remains of five corsac foxes eaten by wolves were found. Wolves
attack mainly animals weakened by starvation, or eat their dead
bodies. They rarely catch healthy animals if these are near a bur-
row in which they can hide. A pursued corsac fox runs with a
speed of 40—45 and up to 50 km/hour, but it can run thus for only
15—20 minutes. The desert wolf runs with a speed of 40—45 and up
to 55 km/hour, but is more enduring and in open places, can catch
the pursued corsac fox, especially if a group of wolves is hunting.
In warm times [of year], wolves attack corsac foxes rarely, but
even more rarely catch them. Thus in Betpak-Dala, the remains of
corsac foxes were found in 0.8% of the wolf feces collected in

311

456

May—June (864 samples). In 101 stomachs of wolves caught in
November—December in the same desert, corsac fox remains were
found in only one. Wolves also destroy corsac fox litters, tearing
apart their burrows (P.T. Kryzhinovskii). However, the attempt to
excavate the burrow does not always meet with success.

The red fox also belongs among the enemies of the corsac fox.
It destroys the corsac fox in winter when it suffers from food
shortages. Cases of corsac foxes hunted by red foxes and car-
casses of corsac consumed by red foxes are known in Kustanai,
Severo-Kazakhstan, Pavlodar, Kokchetav and Tselinograd districts,
i.e. in steppes and semideserts. Red fox attacks on corsac foxes
were observed particularly often in the heavy-snow winters of 1947/
48, 1948/49, 1950/51 and 1953/54. As a rule, hunters found corsac
foxes killed by red foxes in March. With absence of food, red foxes
attack corsac foxes earlier—in November and December (О.М.
Boiko). In summer, red foxes destroyed den burrows of corsac
foxes and killed the young. This was observed in the Oktyabr’sk
region of Severo-Kazakhstan district and the Kellerovsk region of
Kokchetav district (S.G. Chmut and D.P. Fomenko). Decrease in
corsac fox numbers in the semideserts and steppes of Kazakhstan
was related to the rapid growth of the red fox population, crowding
out and destroying the corsac foxes, in the same way as the wolf.
With abundant food, red fox and corsac foxes coexist with one
another. They were observed together at carcasses of fallen live-
stock and shot saigas (Sludskii, 1930).

The corsac fox suffers also from shepherds and stray dogs.
During the driving of livestock to and from summer pastures, thou-
sands of hungry dogs, which harass and kill all animals and birds
which they encounter on their way, go with the herds and flocks
(Sludskii, 1961). Particularly dangerous are the Asiatic borzois—the
taza and vyborzka—which catch up comparatively easily with corsac
foxes. Dogs also destroy the burrows of this animal and wipe out
the corsac pups.

The golden eagle (Aquila chrysaétos) often attacks red foxes
and with training successfully hunts corsac foxes. However, its
influence on corsac fox populations is negligible. The steppe eagle
[Aquila rapax] is a more serious enemy. Remains of corsac fox
were met with several times near the nests of eagle and in their
castings (Nasimovich, 1951). In some places, the steppe eagle is
abundant and may kill a considerable number of corsac foxes.

312

457

Thus, in Kalmykia, in places where saigas give birth to young, 24
to 40 eagles were observed in one field of view (Dal’ et al., 1958).
In rare cases, the upland buzzard (Buteo hemilasius) attacks corsac
pups (Nasimovich, 1951). In Semipalatinsk district, at the end of
winter, remains of three corsac foxes were found eaten by an
eagle owl (Bubo bubo). In winter, in the forest-steppes, steppes
and semideserts, the snowy owl (Nyctea scandiaca) may attack
the corsac fox.

Diseases of corsac fox remain nearly unstudied. It is known
only that they suffer from rabies and carnivore distemper. In 1950
and 1951, a distemper epizootic spread over the whole territory of
Gur’ev and Zapadno-Kazakhstan districts, the western half of
Aktyubinsk district, and adjacent Astrakhan, Volgograd, Saratov
and Orenburg districts. In this vast territory, mass mortality was
simultaneously observed in corsac fox, red fox, wolf, raccoon dog,
steppe polecat and even dogs. Mortality of corsac foxes from
carnivore distemper was noted in 1950 in the Chapaevsk region of
this same area (Sludskui, 1954). Summer mortality of corsac foxes
from an unknown infection occurred in places in the Kurchumsk
region of Vostochno-Kazakhstan district at the end of April 1948,
and in July—September 1951 also in the Leninsk region of Severo-
Kazakhstan district, where separate hunters found up to three fallen
animals. Rodent plague, induced by B. pestis* has not yet been
established in corsac foxes, though several hundred corsac foxes
were examined in the regions where plague is endemic.

In a young corsac fox, caught in the Sovetsk region of Severo-
Kazakhstan district on 27 July 1941, the neck and head were heavily
infected with ticks (species not determined). The following four
species of fleas were noted on animals from central Kazakhstan:
Pulex irritans (large numbers), Ctenocephalides canis (rarely
encountered), Ceratophyllus tristinus (rarely encountered) and
Ctenopthalmus breviatus (rarely encountered). The first two flea
species are specific parasites of carnivores and of the latter two,
one is a parasite of ground squirrels and the other, of voles (Mikulin,
1956).

In southeastern Trans-Baikaliya, for a series of summers in May—
Octrober 6, 413 fleas were collected from 105 corsac foxes between
May and October, from which were identified (in % of total): P.

*Now Yersinia pestis—Sci. Ed.

313

458

irritans L.—70.2; Oropsylla silantiewi W—3.7; Chaetopsylla
homoeus Rh.—S.3; Frontopsylla elata luculenta J. et R—2.0;
Ctenophyllus hirticrus J. et R—11.7; Rhadinopsylla dahurica J.
et R—3.0; Amphalius runatus J. et R—3.2; Ceratophyllus
tesquorum sungaris J—0.1; Neopsylla bidentatiformes W—0.2;

‘Neopsylla pleskei orientalis J. et Arg—0.05; Ophthalmopsylla

praefecta J. et R—0.2; Frontopsylla wagneri J—0O.07;
Pectinoctenus pavlovskii J—0.07; Paraceras flabellum W—0O.01;
Archaeopsylla sinensis J. et R—0.07; Ceratophyllus calcarifer
W.—0.03; Amphipsylla vinogradovi J—0.01.

Fleas not specific for carnivores and which parasitize mainly
rodents in Trans-Baikaliya, occur on the corsac fox and are assign-
able to 13 species and constituted 24.5% [of the total] (Brom et al.,
1948). In southeastern Trans-Baikaliya, the average number of fleas
parasitizing a corsac fox were: in April 3.0, in May 35.3, in June
10.9, in July 37.0, in August 32.4, in September 84.8 and in October
188.9 individuals. In corsac fox burrows, there are many fleas.
Since a great number of fleas parasitize the corsac fox, it is prob-
able that it acts as a vector of fleas infected with plague (Brom et
al., 1948).

The mortality rate among corsac foxes from various factors is
not known. Especially many of them die, apparently, from food
shortage caused by the decreases in rodent populations or their
unavailability due to the deep snow cover, or hard-crusted snow, or
Ice:

In Kazakhstan, many corsac foxes died from starvation in the
winters with prolonged snow cover of 1940/41, 1947/48, 1949/50,
1950/51 and 1953/54. Thus, in January 1949, in the Dzhurunsk
region of Aktyubinsk district, one hunter found 5 corsac and six red
foxes dead from starvation. Starving animals allowed humans to
approach them and they could be caught by hand. In southeastern
Trans-Baikaliya, the deep snow cover is also fatal to corsac fox.
Food shortage in winter in some years is manifested as the most
important factor in corsac fox mortality, leading to a population
decrease.

Population dynamics. In one or another region, often very
broad, the numbers of corsac foxes is subjected to great annual
changes. In years with abundant food, their populations rapidly
grow so long as unfavorable conditions of existence do not occur.
Population growth may be very rapid. In 1930, on Barsa-Kel’mes

459

Fig. 114. Adult corsac fox with throat crushed by red fox near its burrow.
Badkhyz Preserve, southern Turkmeniya. 28 April 1962. Photograph by
A.A. Sludskii.

Island (Aral Sea), where the corsac fox was absent, 3 males and
2 females escaped from cages. In 1933 and 1934, 77 corsac foxes—
the whole population—were caught, which had increased 15 times
in 3—4 years (Sludskii, 1939).

Among the factors reducing the corsac population is a small
number of rodents—voles and pikas in the northern half of the range
and gerbils in the southern desert parts—as well as their unavailabil-
ity due to the deep snow cover (deeper than 15 cm) or the formation
of hard snow crust or ice. At this time, corsac foxes go hungry,
become weakened and die in great numbers from starvation, en-
emies (wolves, red foxes) and diseases. Reduction of the population
usually is carried out in one season.

In the northern Caucasus at the end of the nineteenth century,
the corsac fox was considered a very rare and vanishing species
(Dinnik, 1914; Satunin, 1945). However, in 1921—1925, many of the
animals were found there: One hunter caught 15—30 corsac foxes
each season during that time (Heptner and Formozovy, 1941). Dur-
ing 1926 and especially 1927, the corsac fox population decreased

460

greatly and from 1932 to 1944 their skins were not prepared for
tanning. From the 40’s the corsac population there gradually grew,
and then rapidly increased to 1951. However, in 1952, their number
again fell catastrophically (Vereshchagin, 1959). The cause of the
drop in the corsac fox population in the Cis-Caucasus was strong
dzhut [extensive periods of continuous snow cover] (1949/50, 1950/
Эр 1952/53):

In the steppes and semideserts of Kazakhstan, judging by the
delivery of corsac fox skins to the Irbitsk fair, at the end of the 19th
century, its number fluctuated 15-fold. In the period from 1880 to
1895, it was lowest in 1880, 1890 (in 1881,* only 5 thousand skins
were delivered to the fair and in 1890, 2 thousand; both times were
after the heavy dzhut of 1879/80 and 1888/89). The highest popu-
lations occurred in places in 1883—1886; in 1884, 30 thousand skins
were delivered to the fair, and in the following years, 25 and 15
thousand skins.

In the forest-steppes of western Siberia (Orenburg, Chelyabinsk,
Om’sk and Novosibirsk districts) and in the northern steppe zone
of Kazakhstan (Kustanai, Severo-Kazakhstan, Pavlodar, Semi-
palatinsk and other districts), the corsac fox had nearly disappeared
by 1910 (Korsakov, 1938), and reappeared again in 1916—1917. Its
numbers reached a peak in 1923.

In the dry steppes and semideserts (Ural’sk, Aktyubinsk,
Tselinograd (the former Akmolinsk), Karaganda and other districts),
the corsac fox began to appear in considerable numbers from 1919
on. Before this year, the inhabitants of Tselinograd district did not
know of it. It became especially numerous in 1921 and 1922, but
in 1923, the population began to fall rapidly and by 1924, this animal
had again become rare.

Intensive reproduction of the corsac fox in the beginning of the
20’s was attributed to several factors. In connection with the First
World War, followed by the Civil War, the number of hunters
sharply decreased and the hunting economy nearly stopped around
1920—1921, i.e. for 7—8 years. Many fields were abandoned and in
them there was strong reproduction of rodents. In these same
years in the steppes many livestock, especially horses died. All of
this favored the rapid growth of the corsac fox population, and it
quickly spread in all Kazakhstan and adjacent areas. In these years,

*From context, probably should read 1880—Sci. Ed.

314

461

in Kustanaisk, Severo-Kazakhstan and Pavlodar districts, individual
hunters took 150 to 200 corsac fox skins per season. After the
peak population (1921—1923), 225,445 skins were exported from the
USSR in 1923—1924 and 1924—1925— а record number for the entire
period of existence of the trade (Kaplin, 1960).

The rapid decline in the corsac fox population in the middle of
the 20’s was the result of hunting of carnivores and the ploughing
of fallow and virgin lands. After the end of the Civil War, many
hunters returned to peaceful occupations and the steppes were
flooded with agents of tens of tanning organizations which rapidly.
drove up the price of furs. At this time “everyone who was able
went to the steppe with tools, weapons, shovels and barrels filled
with water and ruthlessly proceeded to kill the little foxes in great
numbers, digging and smoking them out of burrows in groups. This
barbaric ‘hunting’ was practiced especially intensively at the begin-
ning of winter with the first snowfall, when it was easy to find the
corsac burrows through their tracks” (Selevin, 1925). This rapa-
cious taking of corsac fox was also carried out in western Siberia
where, moreover, the animals were pursued on horses with or
without dogs, and poisoned with strychnine. As a result, at the end
of the 20’s, the corsac fox in many places in Kazakhstan and
western Siberia was “almost completely exterminated” (Zverev,
1932). Thus, in Semipalatinsk governance in 1921/22, 1785 corsac
skins were tanned; in 1922/23—8,012; in 1923/24—19,344; in 1924/
25—2,312; in 1925/26—S03; in 1926/27—345; and in 1927/28—only
165 (Selevin, 1925, 1930; V.E., 1928). Overall, in four years, the
corsac population had decreased 117 times. In Kazakhstan (as a
whole) in 1924/25, 23,194 skins were tanned; in 1925/26—19,426;
in 1926/27—7,934; in 1927/28—4,024; and in 1928/29—505 (Gross-
man and Kogan, 1932). In the following years, its harvest was
prohibited.

Mass reproduction of the corsac fox, which was observed in
the 20’s of the present century occurred more or less simulta-
neously over a vast territory. A similar picture of reproduction in
the 20’s and subsequent decrease in their population was observed
in the Cis-Caucasus, lower Volga region, forest-steppes of western
Siberia, and Kazakhstan. An analogous picture was observed in
some places in Turkmeniya. Thus, at the beginning of the 20’s on
the plains at the foot of the Kopet-Dag, corsac fox almost com-
pletely disappeared as a result of the application of strychnine. At

462

one poisoned carcass, as many as 13—14 corsac foxes were picked
up in a night (V.G. Heptner).

In the steppes and semideserts of Kazakhstan, the corsac fox
again began to appear in the middle of the 30’s. Thus, in Kustanai
district in the northern regions, it appeared in 1935, but was still
absent in the Semiozernii and other regions, where it appeared in
small numbers (Naurzumsk preserve) in 1936. Only in the heavy-
snow winter of 1940/41 did corsac foxes here become numerous,
migrating from neighboring regions. A new population peak (judging
by tanned skins) in northern Kazakhstan was in 1938/39. In Kzyl-
Orda and Chimkent districts, the corsac fox was especially numerous
in 1934 and 1935.

The rise in the corsac fox population in the 30’s in Kazakhstan
stopped in the winter 1939/40 when, in many areas, the animals
suffered from a sharp food insufficiency. After this winter, the
corsac population in the north of Kazakhstan, judging by tanned
skins, decreased nearly two times. It continued decreasing through-
out the whole of Kazakhstan after the severe heavy-snow winter
of 1940/41. But in 1944, nearly everywhere in Kazakhstan, the
number of corsac foxes again increased and they were particularly
numerous in the dry steppes and semideserts of Kokchetav,
Tselinograd and Karaganda districts. At the beginning of the 40’s,
the corsac fox population also began to increase in the forest-
steppes of western Siberia, and it appeared in regions where they
had long been absent (Makushinsk region of Kurgansk district,
Sargatsk region of Omsk district and others).

In February—March 1948, in the northern and central parts of
Kazakhstan, corsac foxes died in great numbers from starvation
(voles were few). The winter of 1948/49 had exceptionally deep
snow, and again many animals died from hunger and enemies; the
corsac population greatly decreased (Sludskii, 1953). In this winter,
corsac foxes appeared in significant numbers in the steppes and
forest-steppes of Orenburg district (Darkshevich, 1950). In the 50’s
of this century in the northern steppes of Kazakhstan, an increase
in the corsac fox population was observed in 1952, but they were
especially numerous in 1956. In 1955/56, corsac foxes were also
numerous in the steppes of Altai territory. After the severe winters
of 1948/49 and especially 1950/51, corsac foxes appeared in rela-
tively high numbers in the northern foothills of the Tien Shan in
Kirgiziya and in southeastern Kazakhstan.

316

463

Subsequent significant decrease in corsac populations in the
northern, central and southern regions (Betpak-Dala desert),
induced by the absence of the sagebrush voles and other rodents,
took place in the winter of 1959/60. In the 1960/61 season, there
were few corsac foxes anywhere. In Kazakhstan as a whole,
tanned corsac skins fluctuate approximately 15-fold.

In the deserts of the southern type (ephemeral), judging by the
data on tanned furs, the highest population of corsac fox between
1950 and 1959 was in Uzbekistan in 1950 and 1956—1959, and in
Turkmeniya during the time from 1948 to 1959—in 1952 and 1956;
and the lowest—in Uzbekistan in 1955 after the severe dzhut of
1953—1954, and in Turkmeniya, in 1948 and 1953. In the southern
deserts, the amplitude of population fluctuations of corsac fox is
less than in the steppes and semideserts. In Uzbekistan, tanned
skins of this animal varied only 1.5 annually, and in Turkmeniya, 3.3
times.

In the Daurian steppes of southeastern Trans-Baikaliya, the
corsac fox population varied 18-fold over nine years (from 1940 to
1948) (Yurgenson, 1955). Judging from tanning, the highest popu-
lation of corsac foxes here was in 1946/47 and in 1955. The main
reason for this population growth was migration from the Mongolian
Republic. In separate areas of the steppes, the Trans-Baikaliya
population of corsac foxes changed under the influence of local
factors (N.V. Nekipelov): epizootics among tarbagan marmots, their
extermination by the antiplague organization, mass reproduction of
Brandt’s voles and Daurian pikas favoring the growth of the food
base of the corsac fox, causing increase in its population.

On the whole, the number of corsac foxes in the territory of the
USSR changes annually by 10 up to even 100 times. The most
considerable change in corsac fox populations is found in the
forest-steppes and steppes, where with unfavorable conditions, they
completely disappear over vast areas for several years. Populations
are more stable in the southern (ephemeral) deserts. Before a
population peaks, it increases for several years. The high population
is maintained for 1—2 years, and then decreases sharply. True pe-
riodicity is not observed in variations of corsac fox populations.
During the last 20 years, increases have followed after 24 years,
usually after three years. The amount of tanned skins also fluctu-
ates strongly.

Field characteristics. This animal is very similar to the red
fox, but is two times smaller in size. In contrast to red fox, the fur

464

Fig. 115. Tracks of corsac fox. Left—track of Kazakhstan corsac fox,

V.c. corsac L. (quite large form), on the snow in the vicinity of Dzhanybek

(left bank of Volga), 20 October 1951. Right—track of Turkmentyan corsac

fox, Г. с. turcmenicus Ogn. (small form), on wet sand. Karakum desert 100

km north of Ashkhabad, 21 January 1948. Sketch by A.N. Formozov, about
2/3 of the natural size.

color is uniform reddish gray. The tail is fluffy and long as in red
fox and of one color, but does not have the white tip characteristic
of the red fox. Ears are reddish-brown externally and not black as
in red fox.

Tracks proceed directly—in a line. It steps precisely, track in
track. The distance between tracks is 6 to 8 cm. The size of the
footprint in the dust is 4 x 3 cm. At the entrance to the burrow, it
levels the ground. Near the burrow there are always many feces—
small sausages made up of mouse-like rodent hairs. As a rule, it is
met with singly.

In the case of the appearance of danger, for example, when it
notices a human near its burrow, the corsac “barks”. The agitated
call of this animal is similar to the bark of a red fox, but at the same
time is somewhat reminiscent of the mewing of cat and sounds like
“m-yaaa’’. A disturbed corsac fox also gives another call: “un! un!
un!”. The cry “m-yaaa!” can be also heard at night in winter,

316

465

Fig. 116. Track of corsac fox on snow. Betpak-Dala desert. Kazakhstan.
15 December 1956. Photograph A.A. Sludskii.

when the animals feed on carrion, and also during the time of rut.
Young animals at the age of 1.5 to 2 months, sensing danger, give
а cry “vyau! vyau!” (A.S.)

Practical Significance

Corsac fox is a fur-bearing animal, but its fur is of low value,
because it has quite coarse guard hairs. Previously its fur was more
important. Harvest of this animal has been practiced for a very long
time—in Kazakhstan, for example, since the Bronze Age (“Andronov
culture’) and probably earlier. Later, corsac fox skins were used
for a long time for payment of tribute. Thus, the nomads who
inhabited Kazakhstan under Mongol subjugation in the 13th century
were obliged to pay a tribute of furs, which included corsac fox
skins. At the beginning of the 18th century the Tobol’ and Barabin
Tatars delivered corsac fox skins as tribute (Kirikov, 1959, 1960).

From the ends of the 17th and 18th centuries, corsac fox skins
were a subject of brisk trade between Kazakhs and Russians.

SUT

466

“Kirghiz-Kaisaks caught corsac foxes in the Zayaitsk steppe with
golden eagle and dogs, and overtaking them on horses, killed with
whips a multitude and they exchanged the whole lot, from 40 to 50
thousand, with Russians at Troitsk fort and in Orenburg. According
to the Orenburg tariff, they were placed in value at 40 kopeks
each.' The Kirghiz, in their trade and exchange between each
other, used them almost like money and the price of their goods
was estimated according to the price of corsac foxes” (Rychkov,
1762; analogous information concerning the trade in corsac fox
skins in Orenburg is contained in Pallas, 1773). At Yamyshev fort
on the Irtysh and in Semipalatinsk, corsac fox skins were brought
from Dzhungaria (which then included southeastern Kazakhstan)
(Struve and Potanin, 1867). On the importance of corsac fox skins
as barter, P.S. Pallas (1773) wrote that “As they (Kirghiz-Kaisaks;
A.S.) had no money themselves, they evaluated everything with
reference to horses and sheep, and instead of small coins, used
wolf and corsac fox skins”. Harvest of corsac foxes was also
developed in the 18th century in the forest-steppes of western
Siberia (Pallas, 1786). In Kazakhstan, corsac fox skins were of
great economic importance in the 19th century also. From 1857 to
1861*, from 5,213 to 14,546 corsac fox skins were exported annu-
ally from the Kirghiz steppe through the Petropavlovsk,
Presnogor’kovsk, Omsk and Koryakovsk railway districts alone
(Krasovskii, 1868).

In the 18th and 19th centuries, corsac fox trade was also de-
veloped in Trans-Baikaliya, where in the middle of the past century
corsac fox skins were bought “from the first hand” for 50—60
silver kopeks per skin (Cherkasov, 1884) and exported to China,
where they were in great demand. In the 60’s of the 19th century,
a total 40-50 thousand skins were bought each year in Russia
(Kaplin, 1960).

The size of the corsac fox catch in Kazakhstan and Middle
Asia at the end of the 19th century can partly be judged by the
delivery of skins to the Irbitsk fair (they were not sent to others).
Sent (in thousands of skins) in 1881—5; 1884—30; 1885—25; 1886—
15; 1887—5; 1888—15; 1889—4.5; 1890—6; 1891-—2; 1892—6; 1893—6;
18948; 1895—5. By the end of the 19th century, the corsac fox
trade was mainly harvested in the Aralo-Caspian lowland in the

БАГ that time the price of a sheep was 30 to 40 kopeks.
*In Russian original, 1961, a /apsus—Sci. Ed.

318

467

amount of “more than ten” thousand (Silant’ev, 1898; Turkin and
Satunin, 1900).

In 1923/24, in the USSR, 135.7 thousand skins of this animal
were tanned; in 1925/26—42.9; in 1927/28—12.2; in 1928/29—9,113
thousand. Thereafter, in the majority of regions, harvest was pro-
hibited for a series of years. In 1940, 12 thousand were again
tanned; in 1946—20.4; in 1951—21.8; in 1955—23.8 and in 1956—
28.3 (Kogan, 1931; Kaplin, 1960).

In the beginning of the 20th century, about 15 thousand skins
of corsac fox were imported annually from Mongolia to Kalgan
[China] (Bogolepov and Sobolev, 1911), and in 1927—10,200; 1928—
18,200; 1929—16,600; 1930—3,800 and 193 1—24,000 (Blokhin, 1935).
In this country, not less than 15—20% of the skins obtained remain
in the hands of hunters (Bannikov, 1954).

In the late 20’s and early 30’s of the current century, the world
yield of corsac fox skins ranged from 26 to 50 thousand, of which
fewer than half were taken in the USSR, and more than 50% in the
Mongolian Republic. Before 1938, the majority of corsac fox skins
were exported from the USSR to England, USA, France and Ger-
many. For 1923/24 and 1924/25, 225,445 skins were exported abroad
for the sum of 1,783,107 rubles (Kaplin, 1960). The demand for
corsac skins abroad increased greatly when in 1922/23, a method
was discovered to dye them in various colors while preserving the
silvery tips of the guard hairs.

In the 30’s, as a result of the severe drop in populations of this
animal, tanning of its skins greatly decreased, and together with this
exports declined. After the end of the Second World War, the
reduction in demand for long-haired furs influenced the price of
corsac skins—they were greatly reduced, as a result of which their
export in 1951—1958 was no longer profitable. On the eve of First
World War, a corsac fox skin brought 1.2 dollars on the external
market, in 1923/25—3.85 and in 1943—5.80 for a skin of first sort
of the Petropavlovsk type. In the following years, the price de-
clined, and then began to rise again (Kaplin, 1960).

Earlier, in Kazakhstan and western Siberia, corsac foxes were
caught in jaw traps, coursed with borzoi dogs and golden eagles,
chased down on horseback and killed with whips and soila (long
sticks), dug and smoked out of the burrows with smoke from burn-
ing sulfur or grass, or drowned with water. Lastly, they were poisoned
with strychnine, nuxvomica, and other local poisons. In Trans-

468

Baikaliya, they were caught with jaw traps, “bashmak”’, cherkan*
and other traps which were set at the entrance to the burrow. At
the present time, the main mass of the animals is caught with No.
3 and No. 5 jaw traps and they are rarely coursed with dogs.

The corsac fox is not a significant threat to small-animal hus-
bandry. It usually causes little harm to the hunting industry, especially
in the period of mass reproduction of mouse-like rodents, on which
it mainly feeds. In those years with low populations of small ro-
dents, does it sometimes attack large birds and hares. When
populations are high, it may hunt the yellow ground squirrels and
marmots; therefore in regions where these animals live, the number
of corsac fox ought to be limited. In some regions, the corsac fox
damages the hunting industry, eating animals that have fallen into
traps, or dragging them off. Thus, in the Esil’sk region of Severo-
Kazakhstan district, in the season of 1936/37, corsac foxes ruined
300 polecats which were taken in traps (Sludskii, 1939).

In a series of republics and districts, the corsac fox is protected
as a valuable fur-bearer. Thus, it is completely forbidden to dig
them out of burrows, smoke them out or drowned them. In the
majority of republics, hunting this animal is only permitted in the
season when it has prime fur—usually in November—March. In other
times of year, the season is closed.

The Standard of Fur Raw Materials divides the untanned skin
of corsac fox into four types according to the region where they
were obtained and quality of the fur: Petropavlovsk, Kazakhstan,
Astrakhan and Tashkent, of which the first type is the best and the
latter is the poorest.

In periods when populations of this animal fell catastrophically,
harvest was completely prohibited for many years over large ter-
ritories. For example, in the majority of the regions of Kazakhstan,
hunting of corsac foxes was prohibited from 1928 to 1938.

Hunting was permitted only in seven regions where a ground
squirrel-gerbil industry existed (Irgizsk, Turgaisk, Karsakpaisk and
others). Harvest of corsac fox was prohibited until 1939—1942 in
districts north of Kazakhstan, for example, in Orenburg district
from 1932 to 1939. This prohibition together with other factors
favored the restoration of the corsac fox population. In the gerbil,
marmot and other game industries, corsac fox populations ought to
be restricted, without, however, destroying them completely. (A.S.)

*Local names of different trap types—Sci. Ed.

319

469
Fox, Vixen, Red Fox'®
Vulpes vulpes Linnaeus, 1758

1758. Canis vulpes. Linnaeus. Syst. Naturae, ed. X, 1, р. 40.
Uppsala, Sweden.

1758. Canis alopex. Linnaeus. Ibidem, p. 40. Uppsala, Sweden.

1777. Canis caragan.* Erxleben. Syst. Regni Anim. Mammalia,
p. 566. Area of the lower Irgiz and Turgai [rivers] north of
the Aral Sea."

1789. Canis crucigera. Bechstein. Gemeinn. Nat. Deutschlands,
1, p. 250. Thuringia.

1811. Canis melanotus. Pallas. Zoographia Rosso-Asiat., 1, p. 44.
Kazakhstan south of Orenburg (V.H.).

1816. Vulpes vulgaris. Oken. Lehrb. d. Naturg., 3, Th. 2, р. 1034.

1836. Canis vulpes montana. Pearson. Journ. Asiat. Soc. Bengal,
5, p. 313. Himalaya.

1843. Vulpes flavescens. Gray. Ann. Mag. Nat. Hist., 11, p. 118.
Northern Iran.

1854. Vulpes pusilla. Blyth. Ibidem, p. 729. Salt range, Punjab.

1854. Vulpes leucopus. Blyth. Ibidem, p. 729. Mal’tan (Multan),
Punjab.

1875. (Canis vulpes) var. beringiana. Middendorf. Ubersicht d.
Natur Nord- und Ost-Sibiriens, 4, Th.2, 1 Lief. p. 990. Asi-
atic coast of Bering Strait (V.H.) (“an den West-als an den
Ost-Kusten des Berings-Armes’).

66]

6Нищег$ often call the male “хоум”.

"ЕгеБеп (1777) stated that this form inhabited the “Kalmytsk and Kirgiz
deserts”, i.e. steppes, referring to it in two places in Vol. | of Pallas “Reise”
(1811). The first reference (pp. 199—200) regarding the area south of the Samara
river, is not precisely. determined and may not require attention. The second (р.
234) refers to the area south of Orenburg where “the Kirgizes bring the steppe
foxes (karagan) which do not differ in color from wolves” to be sold in Orenburg
(and hence the diagnosis of Erxleben —‘“colore griseo-lupino”). The above men-
tioned type locality of the subspecies represents a more precise definition to
references based on the usually cited type localities (“Kalmytsk and Kirghizian
steppes”, “Kirghizian steppes”, “Kazakhstan”) which are extremely broad.
Karagans are, apparently, quite characteristic of the suggested limits of the type
locality. Erxleben (1777) considered the karagan as “а doubtful animal” (animal
subobscurum). Later, Pallas (1811) considered it to have the status of a separate
species differentiated from the red fox.

*Sometimes spelled kKaragan—Sci. Ed.

470

1902.

1903.

1905.

1907.

1907.
ТО.

HOW:

9»:

1914.

1922.

320 1922.

Vulpes vulpes splendens. Thomas. Ann. Mag. Nat. Hist.,
10, p. 489. Astrabad, northern Iran.

Vulpes anadyrensis. J. Allen. Bull. Am. Mus. Nat. Hist.,
19, p. 167. Markovo on the Anadyr [river].

Vulpes alpherakyi. Satunin. Izv. Kavkazsk. Muzeya, 2,
No. 1, p. 146. Geok-Tapa. Areshsk Co., Elizavetopol’sk Gub.
[governance].

Vulpes kurdistanica. Satunin. Ibidem, p. 48. Gel’sk depres:
sion, form. Karssk district, Turkey.

Vulpes vulpes tarimensis. Matschie. Wiss. Ergebn. Filchner
Exped. nach China, 10, T.1, p. 164. Tarim.

Vulpes vulpes tschiliensis. Matschie. Ibidem, p. 169. Beijing.
Vulpes vulpes septentrionalis. Brass. Aus. dem Reiche
der Pelze, p. 454. Norway.

Vulpes kamtschadensis. Brass. Ibidem, р. 456. Kamchatka.
Vulpes vulpes flavescens var. cinerascens. Birula.
Ezhegodn. Zool. Muzeya Akad. nauk, 17, p. 254. Eastern
Khorassan, Iran.

Vulpes alopex var. caucasica. Dinnik. Zapiski Kavk. otd.
Russk. geogr. Obshch., 26. Zveri Kavkaza [Animals of the
Caucasus], 2, p. 449. Vladikavkaz (after Ognev, 1926).
Vulpes vulpes krymea montana*. Brauner. Zapiski
Novoross. Obshch. estestvoisp., 40, p. 15. Montane Crimea.
Vulpes vulpes crymensis. Brauner. Ibidem, p. 36. Montane
Crimea.

Vulpes vulpes crucigera n. stepensis. Brauner. Ibidem,
p. 15. Steppes between the Dnepr and Don [rivers], more
exactly—Askaniya-Nova (V.H.).'8

Vulpes alopex var. kamtschatica. Dybowski. Archiw Tow.
Nauk. Lwowe, 1, р. 350. Kamchatka. Nomen nudum.
Vulpes alopex var. sibiricus. Dybowski. Ibidem, p. 350.
Nomen nudum.

Vulpes alopex var. ussuriensis. Dybowski. Ibidem, p. 350.
Nomen nudum.

Vulpes vulpes jacutensis. Ognev. Biologicheskie izvestiya,
1, р. 116. Vic. of Yakutsk.

*Sometimes spelled krimeamontana—Sci. Ed.
8Vicinity of Kherson, after Ognev, 1926.

471

1923. Vulpes huli. Sowerby.* Naturalist in Manchuria, 2, p. 44.
Manchuria.

1924. Vulpes vulpes crucigera diluta. Ognev et Vorobiev. Fauna
pozvonochn. Voron. gub. p. 102, Rocky steppe. Bobrovsk
u[ezd] [county], Voronezhsk. Gub. [governance].

1924. Vulpes vulpes schrenkii. Kishida. Mon. Jap. Mammals,
p. 41. Sakhalin.

1924. Vulpes vulpes splendidissima. Kishida. Ibidem, p. 47.
Northern and middle Kuril Islands.

1926. Vulpes vulpes alticola Ognev. Uchen. zap. Severo-kavk.
inst. Кгаеуедетуа, 1, р. 52. Lake Gokcha (Sevan), Armenia.

1926”. Vulpes vulpes karagan natio ferganensis. Ognev. Annales
Musei Nat. Hungarici, 23, p. 222. Osh, Fergana.

1926. Vulpes vulpes karagan natio pamirensis Ognev. Ibidem,
p. 222. Near Pamir Post, Pamirs.

1926. Vulpes vulpes ochroxantha Ognev. Ibidem, p. 225. Aksai,
Tien Shan.

1926. Vulpes vulpes tobolica Ognev. Ibidem, р. 227. Obdorsk,
lower Ob’ [river].

1926. Vulpes dolichocrania Ognev. Ibidem, p. 232. Sidemi in
southern Ussuri territory.

1929. Vulpes vulpes kiyomassai Kishida et Mori. Lansania, 1,
p. 82. Northeastern Korea.

1931. Vulpes vulpes daurica Ognev. Zveri Vost. Evropy i Sev.
А$п, 2, р. 331. Kharagoi, 45 km west of Troitskosavsk
(Kyakhta), Trans-Baikaliya (V.H.).

Diagnosis

Measurements large—body length more than 65 cm, greatest length
of skull more than 115 mm. General color from light-reddish to
grayish-ochre, brown; dorsal side of ear black, tip of tail white
(V.H.).

*In original text, misspelled “Sowerdy’—Sci. Ed.
‘Issued in the beginning of 1927 (April; V.H.).

321

472
Description

The red fox is a very well-formed, beautiful animal with an elon-
gated body on relatively short limbs. The tail is long (longer than
half the body length), fluffy, and in the standing animal reaches the
ground. The muzzle is quite long and pointed.

In its short summer coat, the red fox appears lean with rela-
tively long limbs, with a thin elongated body, large head and large
erect ears broad at the base and pointed at the tips. Northern foxes,
in their fluffy winter coat, look stocky with short limbs, massive
body and a proportionally moderate-size head. The ears, the basal
portion of which are hidden in the fur, are more proportional [ap-
pear smaller], the tail is especially long and fluffy, its overall length
with hairs is nearly equal to the body length.” Southern forms of
red foxes, in which the winter fur is relatively short, appear in
winter as longer legged and “leaner”, their ears are larger—taller,
and broader at the base.

Pupil of the eye is oval, and vertically oriented. There are three
pairs of teats.

The winter pelage of the red fox is dense, soft, silky and rela-
tively long. However, in connection with its very wide distribution
and its occurrence under extremely different climatic conditions—
from tundra to hot deserts—the geographical variation in this character
is very great. In the northern forms, the fur is very long, dense and
fluffy, while in the southern—much shorter, sparser and coarser
(see section on “Geographic Variation”).

Still more variable is color of the winter fur of the red fox.?!
This [variation] is especially complicated and great because, on a
level with the well expressed geographic variation is wide individual
variation. Besides form and amplitude of the latter, variation also is
observed in types of morphisms. The picture is further complicated
in that in various subspecies, not only is the extent of natural vari-
ability not the same, but also various color phases and types are
encountered in various percentage ratios or are completely absent.
For the latitude of variation in color and quality of fur, this species

?°Hunters very neatly call it “tube”.
2'Color variation in summer fur was not studied, but judged by everything,
it is less.

473

SSS Ee

Fig. 117. Red fox, Vulpes vulpes L. Sketch Бу А.М. Komarov.

is divided into 41 sorts, most of which are in our animals. Within
the sorts, they are separated into different color types. In addition,
parallelism in geographic and individual variation is well exhibited in
red foxes. Therefore, the red fox is one of the most variable in
color of the species of our fauna. Notwithstanding that, variation in
this species is realized through consistent geographic rules.

Two main geographic color types exist—northern red fox and
the southern gray desert fox. To each of these types several sub-
species belong. The extreme forms of both color types differ greatly
from each other (Kamchatkan red fox—Turkmeniyan red fox), but
they are connected to each other by a chain of gradual transitions
through the characters of individual races, mainly those which occupy
intermediate geographic positions, as well as through the individual
variations within the limits of some subspecies which are mixed, in
a geographic and systematic sense. The pelage of middle Russian
and Turkmeniyan red foxes serve as examples of the geographic
coloration of the first type.

Winter pelage of the middle Russian (around Moscow) red
fox. General color bright reddish-rusty with yellowish tint. Along
the spine, weak, diffuse pattern of many brown-reddish-chestnut

322

474

hairs noticeable; from this area, two diffuse stripes pass down-
wards to shoulder-blade. Together with spinal stripe, they form a
striking pattern resembling a cross. In the posterior part of the back
there are often some white or white-tipped hairs, giving the color
in this region a mottled silvery appearance. The sides of the body
are paler than the back; the side of neck is similarly colored. Chin,
lower lips, throat and anterior part of chest white. The remaining
lower surface of body dark, brown or reddish; sometimes a white
stripe extends from chest to anterior part of abdomen. Greater part
of groin dirty white, sometimes almost white, usually with one or
another intensive reddish tint. Posterior part of thighs light and
whitish. Sometimes, the color of the belly and inguinal region does
not differ from that of the sides or even the back. Upper parts of
limbs rusty-reddish. Anterior side of forelimbs, including forepaws,
and paws of hind limbs, black anteriorly.

Anterior part of muzzle, forehead, vicinity of eyes, area be-
tween ears, and upper neck bright brownish-rusty-red. Upper lips
white. On sides of muzzle anterior to eyes, no dark field, or only
a slight darkening (browning) in form of a spot or stripe, passing
from eye to region of vibrissae. Ears, from behind entirely black, or
only the base has a brownish-reddish color similar in color to top
of head. Inner surface of ear covered with whitish hairs.

Tail dorsally brownish-reddish, but less bright than back and
side. Along sides and ventrally, it is pale gray with straw-colored
tint. Many of the hairs, especially dorsally, have dark tips and hence
a more or less intensive, blackish frosting. Sometimes, tail is inten-
sively covered with black, and reddish tones are not developed.
Along tail dorsally, slightly behind the base, usually a dark spot
located in region of supracaudal gland.”* Extreme tip of tail white,
although size of this “flower” is variable.

Foxes of the type described are, in hunting terminology, usually
called “belodushka” [white breasted] or “berezovik” [brown
mushroom]. This is a certain modest type of fluctuating variability:
animals duller and paler in their general tone are encountered, as

2? п a freshly killed animal, this gland, especially in freezing weather, gives
off the odor of violet. This odor is not strong and rapidly disappears especially
in a dead animal taken into a warm location. This explains why the presence of
the “violet” odor is sometimes negated.

*Also called violaceous gland—Sci. Ed.

323

475

are others that are brighter. Especially bright and color-saturated
animals are called “ognevka’™ [fire].

Particular variations in belodushka lie in the distribution and
greater or lesser intensity of the black color on the limbs, in red-
dening of the white area of the thoracic region and lightening
(whitening) of the belly and inguinal regions, etc. Sometimes, the
cross on the back is slightly darker than that described, and is
revealed more clearly.

The other color types of Middle Russian foxes generally rep-
resent stages on the course toward melanism. These mutant color
phase types are often, apparently, hybrids of the normal color type
with melanistic mutants. Melanism in the form of a light admixture
of black pigment appears also in foxes that are in all respects true
“red” type. Sometimes, it is particularly obvious—tt is the “medio-
cre” of furriers. Three types of true, more or less sharply separated,
melanists are differentiated.

Sivodushka [gray breasted]. “Rump and spine brown or gray
color with more or less marked zonation of guard hairs; light bands
of these hairs not white, but yellowish. Underfur on rump and spine
gray, without light tips. Cross on shoulders brown, rusty-brown or
brownish-reddish. Part of fur along sides of this cross are reddish
or yellow color. Sides with large admixture of yellow or reddish
guard hairs. Chest and belly dark brown, sometimes with white spot
in center of chest. Limbs brown”.

Krestovka [cross fox]. “Fur has darker coloration. Rump and
posterior part of spine dark brown or dark gray in color with more
or less well developed silvery color (white zones) on guard hairs;
underfur in these parts of the skin dark gray throughout whole
length of hair. Cross on shoulders black or blackish-brown, some-
times light silvery. Parts of fur at corners of this cross are golden
yellow or rusty. Admixture of yellow or reddish hairs usually also
obvious on anterior part of sides of skin. Chest and belly dark
brown (on chest, a white star sometimes appears). Feet and head
brown”’.** In some places, hunters call red foxes, but with a more

This terminology of Middle Russian hunters does not fully coincide with
the terminology of furriers (see below), especially the term “ognevka’’.

*4According to Kuznetsov, 1952. In furriers terms, the chest is called dushka,
the belly—cherev, and the rump—oguzko. The characters mentioned are standard
characteristics.

476

sharply marked cross-like pattern, a “cross fox”; which is, how-
ever, not black or black-brown, but only a somewhat darker tone
than the general fur color. In exactly the same way, sometimes
forms only somewhat darker below are called gray-breasted, but
they actually are only a transition from the red fox to the true
sivodushka, etc.

Chernoburaya [blackish-brown]. The whole skin is blackish-
brown, or black with light brownish tint. Usually, the skin has a
admixture of various amounts of “silver”, i.e. pure white guard
hairs or black guard hairs with a white zone. Reddish hairs are
completely absent, or they are found in small quantities.”

Between all of these color types—from the ognevka [fire] to
the chernoburaya [blackish-brown], there is a more or less com-
plete chain of transitions. Moreover, in one litter may be encountered
both red, and also melanistic, foxes (Hofman, 1856; Middendorf,
1869; Maidel, 1894; Iokhel’son 1898; Sokol’nikov, 1927; Folitarek
and Chirkova, 1930).

Melanism in foxes is, to a significant degree, geographically
localized. Melanists are more often met with in the forest zone and
in the forest-tundra belt, especially in middle and eastern Siberia,
and also in the Caucasus mountains. In Taimyr, in individual years,
sivodushka, krestovka and chernoburaya occurred in up to 20—
30% of the prepared skins. Melanism is manifested also in other
natural zones, being, however, very rare in steppes and deserts
(Pallas, 1788; Kessler, 1850; Eversmann, 1850; Chernai, 1853;
Bogdanov, 1871; Karelin, 1883; Alferaki, 1891). Appearance of
melanists in one or another place changes yearly. In years of fox
abundance, the relative quantity of melanists increases and in the
years with low number, decreases (A.F. Chirkova). In Canada, the
number of melanists found is proportional to the abundance of red
foxes (Hewitt, 1921).

The “black-brown fox” described as is clear from the above is a wild
melanistic form of the Eurasian red fox. That which ordinarily was usually called
black-brown fox or in vulgar form, “chernoburka,” 15 the so-called “chernoser-
ebristaya”™ [silvery black} domestic fox, raised in captivity, the melanistic form of
the American red fox. It is characterized by pure black color with a variable
admixture of silvery (from 25% to 100% of the skin area). Selection of this form
proceeded by eliminating the brown and establishing in the pelage a pure black tone
of the skin. Brown is considered a defect which lowers its value.

324

477

The described dark forms of foxes are distinguished by the
exceptional beauty and uniqueness of the fur and are valued very
highly (due also to their rarity). Usually (this partially depends upon
fashion) krestovka are valued more highly than sivodushka, and
chernoburaya—higher than krestovka.

Depigmentation of the pelage and complete or partial albinism
(if limbs are white, then foxes are “in white stockings”) seldom
appears among foxes (especially complete albinism). Known cases
of albino foxes more often appear in the southern forest zone, but
they are also encountered in other natural zones (Krasheninnikov,
1755; Eversmann, 1850; Rossikov, 1887; A.M. Nikol’skii, 1889;
Slyunin, 1900; Dinnik, 1914; Pomerantsev, 1939; Dvoichenko, 1955;
Korneev, 1956 and others). Albinism, along with several deforma-
tions, usually develops in foxes after years of insufficient food
(A.F. Chirkova). Chromists appear even more rarely, for example,
red foxes without a white “dushka” [breast], etc.

Summer fur in all formed of red foxes is short, sparse and
coarse. Generally, it has the same color as in winter, but is duller,
dirtier and less bright.

The young fox in the first coat has short, dense wool dark
blackish-brown in color. The lower part of the body is somewhat
grayer, but it is almost undifferentiated from the upper part. On the
forehead above the eyes, there is sometimes a frosting of rusty
tone; on the chest, a small white spot often occurs. The extreme
tip of the tail is white. The replacement of this coat by the second
begins very quickly, in which the reddish tone is well developed.
This is the first adult coat. There is no sexual dimorphism in color.

Winter pelage of the gray desert (Turkmeniyan) fox. The
distribution of the areas of various color intensities is generally the
same as described in the red fox, but reddish color tones are ab-
sent, or in place of them, only light sandy yellow. Dull-yellowish or
gray general tone of fur predominates. Head yellowish-creamy or
sandy in color, with more intense color on forehead and upper part;
muzzle whitish. On muzzle, in front of eyes, there is a well defined
dark spot or band passing from eye to region of vibrissae, colored
dark gray, reddish-brown or blackish-brown. Sometimes they are
weakly developed. Greatest part of rear surface of ear pure black
or brownish-black. On upper neck and back, color more intense,
and yellow or brownish tones more evident in it; it extends over the
shoulders also, forming a cross-like figure. On posterior part of

325

478

body, dorsal stripe more strongly speckled with light-whitish and
light-yellow hairs. Sides gray, without a yellow admixture or with
very weak dilution of this tone and in general is brighter than back.
Along whole body, there is a mixture of black or brown guard hairs,
forming evenly spaced light and dark dappling.

Throat white, chest and belly whitish or gray, chest often being
gray. Feet gray without black stripes, or else an ill-defined darken-
ing present. Color of inner parts of feet yellowish. Tail light
gray-whitish dirty tone. On upper surface near base (in region of
supracaudal gland), is found an ill defined blackish or grayish spot.
Mixture of black hairs along whole tail, more abundant on sides and
on lower surface.

The normal (fluctuating) color variation in foxes of this type is
not less, and in some places, even more than in red foxes. For
example, just in the neighborhood of Ashkhabad, seven color types
are well marked (V.G. Heptner). Among them are encountered all
varieties from pure gray and even fairly dark gray animals to those
in which yellow tones are well developed (all skins appear bright
yellow) and, as a really rare exception, foxes with quite bright
yellowish-reddish color. The lower surface of the body may be
white, or sometimes even brown. The cross is also subjected to
variations. Deviations towards reddish color are, however, more
rarely met with than towards grayish.

Variations in the form of sivodushka, krestovka and
chernoburaya among gray foxes are encountered more rarely
than among the red foxes, and in some territories they are com-
pletely absent.

The summer fur is very short, sparse and coarse; it is charac-
terized by predominance of gray tones, but well developed ochre
occurs.

The fox skull is fairly narrow and elongated, with a small brain-
case and relatively narrow, more or less rounded, zygomatic arches.
In the region of the infraorbital foramen, the skull looks somewhat
compressed, and the line of the zygomatic arches passes fairly
abruptly into the outline of the rostral region. Rostral part of skull
quite long and attenuated: distance from posterior margin of
infraorbital foramen to posterior margin of canine alveolus greater
than skull width above canines. Ratio between length of nasal bone
and that of distance from their posterior margin to most prominent
point of occipital region comprises 1 : 1.2—1.5.

479

Line of upper profile not straight—there is an evident depression
in region anterior to preorbital foramen and forms quite convex line
in fronto-parietal region. Braincase relatively narrow and elongated
posteriorly—posterior border extends significantly backwards behind
line uniting articular condyles of lower jaw. In small southern forms,
braincase somewhat larger relatively and more swollen than in
northern forms. Well defined groove runs from level of supraorbital
processes along suture between frontal and nasal bones.

Occipital crest well developed; sagittal crest quite well devel-
oped in some forms (large northern foxes) and in others (small
southern forms)—very weakly. In first, temporal lines pass back
from postorbital processes at acute angle in respect to each other.
They enclose a small, narrow triangular area and pass to arrow-
shaped crest at level of, or immediately behind coronal suture (in
old individuals). In small Middle Asiatic forms, temporal lines run
almost parallel, outlining a quite large area, and then diverge, joining
small arrow-shaped crest near occipital (here, one of the evident
“infantile” characters of the small southern form of fox is shown,
both in development of crests and structure of braincase itself).
Ends of paroccipital processes located only slightly lower than level
of lower margin of auditory foramen. Length of symphysis of lower
jaw 4—5 times shorter than greatest length of entire mandible.

Canines relatively long: when jaws are closed, lower extend
beyond edge of upper alveoli; ends of upper canines (if skull is
viewed from in front) extend below lower border of anterior part
of lower jaw.

Individual variation in skull quite significant not only in general
dimensions, but also in proportions. Skull within limits of species
norms, may be relatively short and wide, or narrow and long (elon-
gated). Description of a separate “species’—V. dolichocrania
(Ognev, 1926)—was based on a specimen with an extreme degree
of dolichocephaly.

Sexual differences in the skull are expressed in smaller average
size of the female skull, and this difference is more pronounced
than in corsac fox. As a group, females have, apparently, a some-
what wider nasal region and hard palate, canines are somewhat
larger and the upper tooth row is longer (Ognev, 1931). However,
these characters require verification. Apparently, females have
somewhat weaker skull sculpturing.

327

480

Age variation in the skull proceeds, in the direction of growth
in the relative size of the facial region and decrease in that of
cranial region and relative decrease in size of braincase. The oldest
animals have skulls with especially elongated facial parts, relatively
small braincase, and well developed temporal lines and crests.
Several proportions and structural characteristics of the skull exhibit
geographic variation (see beyond).

The diploid number of chromosomes is 38.

Dimensions of the fox exhibit significant age, sexual and indi-
vidual variations. Moreover, geographic variation is also great. At
the same time, up to the present there is little accurate information
on body dimensions and weight of the animals; data concerning
skull size are considerably greater.

Body length of adults is 49—90 cm, tail length without terminal
hairs—53—60 cm, ear length—7.7—12.5 cm, length of hind foot is 12—
18.5 cm. Height at shoulder 1$ 35—50 cm.

Weight is 2.2—10 kg. Females are somewhat smaller and lighter
than males; in each locality, their weight 1s approximately 15-20%
less.

Maximum skull length of males is 129—167 mm, females, 128—
159 mm; condylobasal length of skull in males is 125—160 mm,
females, 115—145 mm; zygomatic width in males is 64—89 mm, fe-
males—61—83 mm; cranial width in males is 41—52 mm,
females—41—51 mm; length of upper tooth row in males is 60—70
mm, females, 52-69 mm.’

Systematic Position

Apparently, the red fox must be considered a more specialized
species than the above-described V. cana and V. corsac, as well
as V. bengalensis. In any event, the skull of V. vulpes, even the
smallest forms, possesses fewer infantile features than the skull of
the mentioned species. The great development of the facial part of
the skull must be considered a large specialization in the direction
of carnivory. The large size of V. vulpes may also be considered
as a progressive feature. Characteristically the southernmost forms

Тре figures given relate only to adult animals, but are attributed to the
species as a whole (within the USSR)—data on separate subspecies are given in
the section “Geographic Variation” (V.H.).

481

> и, HB
Ws FF

SASS
м. АИ WM, № “А a
\\ vy! \ NIN Ay `
== Nae \ \ Way
ААА = Мих
\y)! TA
b

C76

326 Fig. 118. Skull of red fox, Vulpes vulpes L.

328

482

of V. vulpes (griffithi, pusilla, from ours—/lavescens), as com-
pared to the northern ones, have more primitive features of the
infantile type—the relative development of the facial region, devel-
opment of crests and several others. In a certain sense, they show
transitional features from V. vulpes to the above-mentioned small
species.

Nevertheless, within the genus Vulpes, s. str. V. vulpes must
not be considered the final link. The Tibetan Г. ferrilata represents
a species which is still more specialised, in the mentioned trends,
than V. vulpes. This is a higher manifestation of the fox type.

The North American red fox is identical with our fox in regards
to species, although the American authors considered it up till now,
a separate species—V. fulva Desm., 1820 (Hall and Kelson, 1959).
Specific identity of both species is apparent not only from their
morphological characteristics, but also their zoogeographic relation-
ships. Both forms can be freely crossed in captivity and yield
normally fertile offspring. Probably, this also occurs in nature. In
any event, the European red foxes brought into the eastern part of
the United States are indistinguishably mixed together?’ (V.H.).

Geographic Distribution

The range of the species is very vast, and occupies the Old World
from the Arctic Ocean southward to North Africa, Arabia, north-
ern India and northern Indochina. In the New World, the range
extends from the north of the continent to the northern shore of the
Gulf of Mexico.

2’The most recent information on this question is as follows: “Red foxes
(Vulpes vulpes) were introduced numerous times from England between 1650 and
1750, and may have become naturalized or crossed with our native red foxes (V.
fulva). The survey did not contribute any new information on the debatable
question of whether present fox populations of the east-central and southeastern
states are entirely native, a mixture of native and introduced animals, or entirely
from introduced animals, as deduced by Gilmore (1946)” (Presnall, 1958).

“Whether the red fox of the eastern United States (much or all of the area
shown, on the distribution map as the range of Vulpes fulva fulva) 1$ a native animal
or instead a stock introduced from Europe is unknown at this writing” (Hall and
Kelson, 1959).

483

Geographic Range in the Soviet Union

This comprises a very large part of the species’ range, and
covers the entire state territory except the very far north.

The northern border includes the whole of the Kola peninsula
together with neighboring islands (Kil’din), Solovets islands, Kanin
peninsula, Kolguev Island, Vaigach and Yuzhnyi islands of Novaya
Zemlya—along the western coast to Gusinaya Zemlya inclusive. In
Yamal, the border passes through the region of crooked-forests
[{krummholz] (forest-tundra), approximately along the Khadyt river.
Farther east, it coincides with the sea coast of the Ob’ Gulf and
extends to the mouth of the Taz. Hence, the line of the border,
apparently bypassing the Gydansk peninsula, suddenly ascends to
the northeast, crosses the mouth of the Yenisei in the northern part
of Brekhovsk islands (red fox normally occurs around Dudinka),
passes on to the lower course of the Pyasina, where it reaches
73° М. Lat. (mouth of its left tributary, the Kura).

In Taimyr, the northern border of the range ascends to the
latitude of Lake Taimyr (about 74° N. Lat.; E.I. Shereshevskii),
whence it descends to the mouth of the Khatanga, including its
large left tributaries, the Balakhna and Gusikha (to 70°30' N. Lat.).
Farther to the east, the fox is met with along the lower Lena
(occurs around Bulun), and in the Lena delta and along the shore
to the east (Tiksi, Cape Bykovsk—72° N. Lat.). From the Yana to
the Indigirka, the border coincides with the coast of the Arctic
Ocean and, apparently, runs even to the Bering Strait itself. The
eastern border of the range consists of the Pacific Ocean. The fox
lives on Karaginsk Island, on all of the Kurils (except the very
smallest), on Sakhalin and Shantar islands; on the Commander is-
lands they are absent.

The border as indicated is approximate to a certain extent. The
northern border of permanent denning of the fox is quite well de-
fined by the northern border of the forest-tundra, although in open
tundra, the animal also reproduces in some places, sometimes even
quite far to the north (Kanin). However, this rarely occurs. At the
same time, the fox penetrates the tundra, especially in the south,
very often. Therefore, it is difficult to determine the northern limits
of the regions of settlement and of transgressions. Foxes pass
especially far to the north along river valleys overgrown with arbo-
rescent and shrubby vegetation far beyond the limits of forest-tundra.

331

484

Apparently, Vaigach and Novaya Zemlya especially represent
regions of transgressions by the fox as well as extreme northern
points in eastern Taimyr. As well as short-distance dispersal, at
some places very distant ones occur. Thus, in Yamal, a fox was
observed in the tundra area, even to Cape Drovyanyi (northern
termination of the peninsula), and in western Taimyr, nearly to
Dikson (Slobodsk Bay). Transgressions of the fox on to the
Lyakhovsk islands are known, particularly to Bolshoi Lyakhovsk, on
the ice of the Chukotsk Sea 100 km from the continent, and even
to Wrangel’ Island. In the latter case, the animal must have trav-
ее not less than 150 km” across the ice. These transgressions
are, of course, extremely rare. Thus, on Bolshoi Lyakhovsk, only 2
foxes were caught in a period of several years. During this same
period, 5,500 Arctic foxes were captured (E.I. Shereshevskii).

Geographic Range outside the Soviet Union

This includes all of Europe, northward to North Cape, on the
west including Ireland, southward to Sardinia, Corsica, Sicily and
Cyprus; in Africa—lower Egypt, the Fayum, Libya, Tunisia, Algeria,
Morocco, Rio-de-Oro and Senegal. In Asia, the range includes
Asia Minor, the entire Arabian Peninsula, Iraq, Iran, Baluchistan,
Afghanistan and all of Central and Eastern Asia, to the south in-
cluding Kashmir, Punjab, Sind, Kutch, Kumaon, the Himalayas
(including Sikkim), south China (Fujian, Yunnan) and Bakbo (Tonkin
Gulf) in North Vietnam, and also the Japanese islands.

In America, the range includes the continent on the north to the
Arctic coast, and the coast of the Bering Sea. The southern border
of the range is complicated. To the east of the 90th meridian, it
reaches the Gulf of Mexico; however, foxes are absent in Florida
and in the eastern parts of Georgia and North and South Carolina.
To the west of the indicated meridian, the fox is absent in a vast
area extending in a north to south belt from Texas, Oklahoma, and
New Mexico in the south to the southern part of Saskatchewan and
Alberta.* It is also absent in a narrow strip along the Pacific coast,
beginning on the Kenai Peninsula, and in parts of California,

8Range according to data of Vrangel’, 1841; Middendorf, 1860, 1869, 1875;
Zhitkov and Buturlin, 1901; Zhitkov, 1904, 1913; S. Naumov, 1931; Ognev, 1931;
Adlerberg, 1935; Kolyushev, 1936; Mineev, 1936; Portenko, 1941; Novikov, 1956;
Heptner, 1936 and others and according to unpublished material of E.I. Shereshevskii.

*The red fox is now present in most of this area—Sci. Ed.

485

Nevada, Utah, Oregon and Idaho. The southernmost limits of fox
distribution in America, starting from the west, lie in California,
Arizona, New Mexico, Texas and Louisiana (from Hall and Kelson,
1959).

On the American Arctic archipelago, there are no red foxes,
but they are found in Newfoundland, islands of the Gulf of Saint
Lawrence, and on the Atlantic coast, and on Kodiak and Saint
Lawrence islands in the west. It is absent on other islands of the
Bering Sea and Pacific Ocean.

The European red fox has been acclimatized in Australia and
inhabits nearly all of that continent, and the European (English)
form was imported into North America (see above) (V.H.).

Geographic Variation

Geographic variation in the red fox is very great. It is greater than
in all of our other carnivores, and greater than usually occurs in
mammals of that size, and with similar biology. This concerns not
only the number of geographic races which it forms, but also the
amplitude of geographic variation in morphology. This is related to
its broad range, and occupation of very diverse natural situations
and entirely variable biotopic relationships. Thus, if the fox of the
northern taiga lives with a winter lasting 7—8 months, with bitter cold
and deep snow, then the animals of southern Turkmentya basically
do not know true snow cover, but live with severe heat and mois-
ture deficit for almost half the year. There are as many sharp
differences in nutrition as well. A large role in the wide geographic
variation of the red fox is played by its extreme general plasticity,
including individual variation. This is greater than in other, just as
widely distributed, species. Thus, individual and geographic varia-
tion in the wolf and other foxes cannot be compared with that of
the red fox.

The picture of the geographic variation in red fox is very com-
plicated and difficult to study. Changes are observed in general
dimensions of body and skull, comparative measurements of some
body parts (ear, tail), several particularities of skull structure (see
above), particularities of pelage (see above), thickness of the skin
(the hide itself) and lastly the color. If in the main all listed catego-
ries of features are “normal”, nonetheless some have individual
variations of significant amplitude, and in the character of color

‘(auido `9’Л) YOu ay} 0} suoIssaissuen Buoy Ajjetsodsa ayy Jo $99814
рие иоцоэлр 94} MOYS змоллу “YSSN ш "1 sadjna sadjn4 ‘хоз dy} JO вэле иоцп91351р dy} JO злэрло4 и1э5еэ рие UJOYVON ‘611 “SIA 658
021

Г ГЕ
АНЯ

Ke \ ae : |
\\

SS
Ce

awe

160 90
pr
ba e
SD
у = > Д
4
e у a)
% Q A

cet и
7.

120 160 180 160

332

488

greatly exceed the “normal” range in other species, even those
which are highly variable in this respect. Practically every geo-
graphically limited, and in a systematic sense, homogeneous
population has some more or less well differentiated types of indi-
vidual variations of the basic (“normal”) color phase. This concerns
the general tone and intensiveness of color, the degree of manifes-
tation of the dorsal stripe and shoulder stripes, foot color, color of
the venter—chest and belly, the character of facial pattern, etc.
These types are characterized by one of the features mentioned or
a combination of them.

In individual variation in color of the different geographic popu-
lations the following particularities exist, important for an
understanding of geographic variation in the species.

1. In various geographic populations (subspecies), the number
of color types differs; more or less variable populations exist. Thus,
in the population in the vicinity of Ashkhabad one may distinguish
7 types of individual color variations (V.G. Heptner). In others, 2,
3, or 4 color types are designated, so on.

2. The portion of each of these type is not identical, and equally
with one, or one or two predominant ones, others are encountered
more rarely, and the remaining are very rare. In general, there is
usually a normal curve of distribution.

3. In various geographic populations, there are different types
of individual variations. Thus, in the north there are no gray foxes,
nor in the south ognevka [fire] (see below). The more distant
geographically the populations are, the more they differ from each
other in the set constituting their color types.

4. In populations, the set of color types which are more or less
the same, may differ fundamentally in the percentage of different
color types contained, and various color types may predominate;
the prevailing color type of one may be rarer, or even rarer in
another [color type]. In this way, each geographic population is
characterized by: (a) its set of types of individual variations and
(b) prevailing type (or types) of color.

5. These relationships become complicated, and general vari-
ability of the population greater, in regions where one defined form
of the population (or subspecies) is contiguous with and mixes with
another (“introgress with one another’), especially if several come
together. This occurs, for example, in regions where vast flat lands

_ are contiguous with mountains (eastern Trans-Caucasus).

489

6. Color type sets are especially varied, the relative percent-
ages are especially complicated, and sharp variations in these
relationships are seen in small areas in those countries where in
relatively small areas, landscape-geographical, zoogeographical and
ecological conditions and relationships change abruptly and are com-
plicated, as for example in the Caucasus (see below, and detailed
material concerning this question in Vereshchagin, 1960).

7. Geographic difference in the distribution of color phases
(sivodushka, krestovka, chernoburaya) and their hybrids with
the “normal” foxes are related to the variations of the “normal”
color type. These phases, despite their scarcity, are found in some
regions more often than in other regions, and in still other regions
they are completely absent.

In the above-mentioned situations, geographic variation in the
fox is manifested especially sharply, especially in color, and with an
amplitude greater than in other characters.

As a result of extensive study of hundreds of thousands of
skins of animals caught annually for the fur trade, 41 “kryazh’*
of the normal color phase of fox are differentiated as regards color,
fur quality and in part, size. In these types, which are characterized
by one and sometimes two prevailing color types, geographic varia-
tion of the species is well manifested. However, there is no full
coincidence between the types and subspecies. In many places,
within the limits of a population belonging to one subspecies, pre-
pared fur is assigned to several types depending upon the properties
of the fur. Thus, in the vicinity of Ashkhabad, foxes are sorted into
2-3 types, and in various parts of Caucasus, where physical con-
ditions are especially complicated, they are sorted into 2, 4, 5 and
even 8 (Vereshchagin, 1960). Therefore, the type is characterized
by both the actual geographic group, and by individual deviations.
Usually only groups of types are coincident with geographical races
(subspecies). A mixture of the types in some places is, apparently,
connected with the fact that here is found an area of mixing and
introgression, of entirely different geographical populations (subspe-
cies) as regards the color. It may also be connected, to a certain
degree, with hybridization.

An attempt is often made, especially by hunters, to relate the
defined color types of foxes in a given limited place to ecological

*Lit., “blocks,” or types—Sci. Ed.

333

490

variability (“reed”, “steppe”, “montane”, “marine”, “tundra”
foxes, etc.). However, there is no foundation for this. Only in some
places in the foothills or the base of the mountains, may montane
and plains foxes actually be differentiated. But this is only in those
cases when the plains are occupied by one subspecies, and the
adjacent montane country—by another, and individuals of both sub-
species are met with in a defined zone. Such a picture exists,
apparently, in places in southern Tadzhikistan, where the plains are
inhabited by V. v. flavescens and in the mountains of Pamir-Alai,
V. у ferganensis is distributed (Flerov, 1935; Chernyshev, 1958).
The described complexity of color variation in the red fox makes
the study of geographic variation in the species very difficult.

A series of described subspecies are unfounded, since they are
separated on the basis of individual deviations. On the other hand,
color characteristics actually present can only be established by a
large series and in an entirely general form according to an aver-
age—the prevailing color type. The establishment of all amplitudes
of variability in this feature is practically impossible at the present
time.

In actuality, all geographically varying characters of red foxes
do not develop irregularly and independently of one another, but are
parallel and geographically regulated. In their development, several
general rules of geographic variation are well revealed.

Northern foxes occupying the forest zone are the largest; also
large are those of the forest-steppe and steppes of the European
part of the Soviet Union, and the northern steppes of the Asiatic
part of our country. Foxes of the southern steppes and semideserts
are smaller, and animals of the extreme south—the deserts of Middle
Asia—are the smallest; foxes of extreme southern Turkmeniya
(Badkhyz) are particularly small. Foxes of southern Afghanistan,
Pakistan and India are still smaller. In the south (Middle Asia and
Trans-Caucasus), montane races of foxes are somewhat larger
than those inhabiting the lowlands.

The average area of a fox skin in Bashkiria (Bashkirian sort)
is 2,908 cm?, that of foxes of the Middle Asian plains (smallest—
Tashkent type)—1766 cm’. The weight of 100 skins of Bashkirian
fox is 40.5 kg on the average; that of Middle Asian plains—27 kg
(Kuznetsov, 1952) (skin weight characterizes length and density of
fur, but mainly general skin dimensions). Weight of Middle Russian
(Moscow) foxes is 4-10 kg (Ognev, 1931); weight of foxes from

334

491

the plains of southern Tadzhikistan (right bank of Amu-Dar’ya) is
2.23.2 kg (Chernyshev, 1958). The large foxes of the forest-steppe
zone of the European part of the Soviet Union have a body length
up to 90 cm, averaging 72 cm; the foxes of southern Tadzhikistan
are only 49-57.5 cm. The weight of a carcass without skin is in the
first case usually 4.4-8 kg, and in the second, averages 2.7 kg
(Chernyshev, 1958; A.F. Chirkova). Foxes of the steppes and
semideserts of Asia and certain other intermediate regions (some
parts of Caucasus) occupy a transitional position between these
two extreme groups.

Skull size also very clearly decreases from north to south, reaching
its minimum on the plains of Middle Asia, and especially in southern
Turkmeniya. Condylobasal length of the skull of foxes of the upper
Pechora (Pechoro-Ilychsk preserve) varies in males (28) from 131.5
to 158.9 mm, resulting in an average of 145.4 mm. Such dimensions,
apparently, have foxes of the entire taiga zone, although larger
individual populations occur. Thus, in the foxes of Kirov district (26),
average condylobasal length of the skull is 149.6 mm. Foxes of
Middle Russia and the northern steppes of the European part of the
Soviet Union (Voronezh, Poltava) are similar to Pechora foxes. The
smallest fox of the plains of Middle Asia (14, Ashkhabad) has a
condylobasal length of adult male of 125.7—139.1 mm, on average 132.4
mm. In foxes from Kushka, this measurement in males is 129.9 mm
and in females, 123.5 mm. Therefore, maximum size of the southern
form is less than the average size of the northern form, and the average
skull size of the smallest race constitutes about 88% of the larger.
Skulls of foxes living still farther south (Indian, Pakistanian and southern
Iranian forms—griffithi, pusilla) is still somewhat smaller. In the inter-
mediate regions, there are populations of transitional character. In the
south (Caucasus, Middle Asia), montane forms of foxes are usually
somewhat larger that those of the plains. Bergman’s rule is well ex-
hibited by the described species.”

Geographically oriented changes in qualitative skull characters
exist. The skull of the northern fox has a relatively smaller cranium,
and strongly developed crests; there is quite well developed sexual
dimorphism in size. Small foxes of Middle Asian plains have a

2Уашез in unpublished material of У.Е. Yakob’. For foxes of the Tadzhikistan
plains, the trend is toward even smaller values (minimum—1 15.1 mm. Chernyshev,
1958), but they belong, however, to specimens that are not fully mature.

335

492

relatively large cranium, and more weakly developed crests (see
above and Fig. 121). On the whole, the skull of the southern fox
possesses infantile features and, in connection with this, sexual
dimorphism is relatively weak.

Variations in body proportions are expressed in that southern
foxes possess a relatively longer tail and longer ears (Allen’s rule).
Montane subspecies in the south (Middle Asia, Caucasus) possess
relatively shorter ears than plains forms at the same latitudes (A.F.
Chirkova). Southern foxes have, apparently, somewhat longer limbs.

The length, fluffiness and softness of the fur varies regularly in
the same manner (from north to south). Northern foxes have dense,
long, soft and silky fur, the southern—sparser, shorter and coarser.
In foxes of the Ob’ lowlands (West Siberia), length of guard hair
on the sacrum (“оп the rump”) on average equals 69 mm, that of
underfur—52 mm. The same indices for foxes of the Middle Asian
plains—48 and 35 mm, which constitute only 70% of the northern.
The coefficient of softness of rump guard hairs in Yakutsk fox
(especially soft hair) is equal to 1.3; while in the coarse-haired
southern foxes (Caucausus), it registers 2.2 (Kuznetsov, 1952)*°. In
the south (Caucasus, Middle Asia), montane subspecies of foxes
have denser, longer and softer fur than plains forms in the same
latitudes.

The thickness of the skin (“hide”) significantly varies geo-
graphically. In northern foxes, it is thin, in foxes of the middle zone,
of moderate thickness, and in southern, thick and rather rough.
Thus, in Yakutsk fox, thickness of the skin ranges between 0.22
and 0.24 mm, with an average of 0.23 mm, while in the Kazakhstan—
from 0.40 to 0.42 mm, with an average of 0.42 mm, 1.е., it is nearly
twice as thick (Kuznetsov, 1952).

Color for all of its variegation and variability, shows completely
regular changes geographically. This is revealed in variation in the
general intensity and saturation of color of the main types, and in
the replacement of these main types when subordinate ones come
to prevail, and similarly in changes of the set of color types them-
selves. Elements of the whole spectrum of color types have an
amplitude from bright red-ginger to gray in some regions, as if
successively displaced from one territory to another. At the same

Coefficient of softness is the ratio of the hair thickness in microns to. its
length in millimeters. The character of the skin can also be judged by skin weight,
as given above.

334

493

Fig. 121. Skull of Middle Russian red fox (V. v. vulpes L—on the left, Kirov

district) and Turkmeniyan red fox (V. у. flavescens Gray, Turkmeniya,

Repetek). Males with similar degree of tooth wear. Sketch by N.N. Kondakov
from material of the Zoological Museum, Moscow University.

time, the role of each color type successively changes from null to
predominance (and vice versa).

The brightest, most saturated reddish-ginger “ognevka” [fiery]
color tones are in red foxes in the northern Far East, including
Kamchatka and Yakutiya. They have almost no bright ripples on
the spine and rump and the sides are brightly colored. On the
anterior side of the foot, there are large, bright black spots. In these
regions, the “ognevka” is evidently prevalent. In all of the remain-
ing forest zone, the bright-reddish color is distributed, but is not the
saturated hue of the “red fox” (as called by furriers). They have
a well defined light, bright ripple on the thighs and in the posterior
part of the spine. Sides are relatively light. On the feet are large
well defined black spots.

In the forest-steppe districts of Siberia, in the steppes of the
European part of the Soviet Union, 1.е. south of the forest zone, and
in the mountains of Middle Asia, lives the “crimson fox”, which is
less intensive colored than the forest one. In them the spine and

494

cross on the shoulder are bright-reddish or rusty-yellow in color, the
sides are sandy-yellow. The dark spots on the feet are small, nar-
row and dark-gray in color.

Farther to the south, in Kazakhstan and Middle Asia, the color
of fox either becomes lighter or grayer. In the semideserts and
deserts live “light’’ foxes in which the spine is a light-sandy color,
usually with a lighter band, the sides are whitish and very light, and
on the feet is a narrow band of dark-gray, and “grays”, in which
the spine and sides are gray in color, sometimes with weakly de-
fined dull-reddish tones along the backbone. The foxes of the
mentioned regions are usually known as karaganka [pea shrub
foxes]. “Grays” represent the extreme degree in loss of reddish
pigment. They predominate in the extreme south of our country.
Between the districts occupied by foxes of the deserts (“light” and
“отау”) and the Asiatic forest steppe lives a population of a more
or less intermediate character. In this way, a successive change in
fox color from “fiery” to “gray’?' runs from the northeast of the
continent to southern Turkmeniya.

A legible picture of pigment loss is also observed in the foxes
of Caucasus, but it is very complicated. Here is encountered the
“red-gray” fox, in which the spine and sides are of gray in color
with a clearly defined rusty band along the backbone, and also the
above-described “gray” fox, several types of red color with yellow
tones; gray and red-gray foxes are mainly native to the plains of the
Trans-Caucasus, and in part the Cis-Caucasus; the remainder are
in the Cis-Caucasus and the montane parts of the country.”

3\Оп a plane with the above-mentioned main types, furriers, in separate
places, differentiate other deviant or transitional types—light-red and dark-red,
light-scarlet, light-yellow, scarlet-yellow, red-scarlet, brick-red, brownish-red,
light-gray foxes, etc. Local hunters, nearly everywhere, in their turn distinguish
various forms, mainly on the basis of color: in addition to above-mentioned
forms, “serebryanka” [silvery], “red” “berezovka” [birch], “belodushka”
(white chest), etc.

Characteristics of the color types of fox were mainly given according to
Kuznetsov, 1952. Therein is presented also detailed data on fur types and their
peculiarities in various regions.

32Рог details on the distribution of the color forms and races of Caucasian
foxes, see Vereshchagin, 1960 (maps).

336

495

As was often shown above, there is known a directional
geographic variation in the phases of color types also (Arestovka,
sivodushka, chernoburaya).

Fur quality also changes noticeably and regularly. The most
fluffy, fine-haired, silky and lustrous furs are possessed by the
foxes of the forest zone, especially of Kamchatka and the adjacent
districts of East Siberia and the northern Far East. In the steppe
and desert zones, the quality of the fox fur is greatly deteriorated—
it becomes short and coarse, losing its luster, silkiness and elasticity,
the hairs are easily matted. Some of these qualities appear in the
open landscapes in the Far North in the tundra zone. On ascending
into the mountains, even in southern latitudes, the quality of the
pelage is again improved as compared with the plains.

In summary, the reality of geographic variation in the red fox
is very nicely characterized by prices in the international fur mar-
ket: ognevka of Kamchatka are 10 times more expensive than
gray Turkmeniyan.

Subspecies of Red Foxes in Our Country

The infraspecific systematics of our foxes is not sufficiently eluci-
dated. This concerns both morphometry and, in particular, color
characteristics. Up to the present, only one attempt at a more or
less full revision of all forms was made by S.I. Ognev (1931, 1935).
This author recognized 20 forms of red fox for the pre-war territory
of the USSR. One of these forms he considered a separate spe-
cies, V. dolichocrania but the other 19 as forms of V. vulpes, of
which two he considered as “tribes” of Г. у. caragan. Since then,
only some remarks were made on separate forms and only very
general remarks were made on all forms (Flerov, 1935; Kuznetsov,
1944; Bobrinskii, Kuznetsov and Kuzyakin, 1944), and a review of
Caucasian foxes (Vereshchagain, 1960).

The number of forms (20) accepted by Ognev (1931) is too
great, but the remarks of Flerov (1935) on the contrary simplify the
picture too much. A solid new review of the question is necessary.
Preliminarily, if one considers only the best characterized forms
(not too “subtle” and generally questionable), 14 to 15 subspecies
can, apparently, be accepted for our country.

Below are given only very short characterizations.

1. Middle Russian fox. V. v. vulpes Linnaeus, 1758 (syn.
septentrionalis).

337

496

Pelage bright, red, but with strongly developed whitish and yellow
ripple on posterior part of back (see above, “Description” section).
Fur long and fluffy or of moderate length and fluffiness.

Measurements large. Body length 70-90 cm, weight 5—10 kg.

Maximum male skull length known is 163.2 mm. Some local
populations within subspecies limits may on average be somewhat
larger (maximum skull length 155.7, condylobasal length 149.6 mm;
Kirov district).

In northern and middle (forest) districts of the European part of
the country southwards to. forest-steppe and eastwards approxi-
mately to the Urals.

Outside the USSR—in Scandinavia and probably Central and
Western Europe.

The Middle Russian fox is usually accepted, following Ognev
(1931), as related to the Middle European V. v. crucigera and
separated from the north Russian, which is supposed to be the
nominal [form]. Foxes of the extreme northern forest zone of the
European part of the Soviet Union, as compared to these near
Moscow, possess a somewhat longer and fluffier fur. However, in
this feature the transition is unnoticeable, the difference not great,
and differences in other characters are absent. According to
craniological characters, foxes of Arkhangel’sk district, the upper
Pechora and Kirov district are completely identical with those near
Moscow (V.E. Yakobi). Therefore the separation of two subspe-
cies in this part of the forest zone of the Soviet Union is not
justified.

The relationship between the form crucigera and the nominal
[form] is not evident even for Western Europe. In our materials it

Table 38. Skull measurements of Middle Russian fox (materials from
Moscow district; V.E. Yakobi)

Data Males Females
М. - Min. Max. Aver. М Min. Мах. Aver.
Greatest length 30 138.3 160.5 151.5 26 138.3 158.8 145.9
Condylobasal 29 1395" 1548 1455 26 1344 1475 14083
length

Length of upper 30 52.2 59:51 55.8127 50.6 58.4 54.4
molar row
Zygomatic width 30 72.8 88.5 79.9 26 73.6 82.7 7:2

Greatest width 30 46.2 SED 486.26 45.1 50.5 48.1

“AVSIOAIUL] MODSOP “WNasN] [BOISO]OOZ sy} JO зремэуеи WO}
AOyepuoy ‘мм Aq Зипитеа (peqeyyxysy Jesu) Ае10 suaasaanyf ‘a “4 ‘хоу URAIUOWYIN | (39113$1р ysuIqnAyyy “uoIsoI Ysuluqe |) УХА и7280402 “AY
‘едиеЗелеу {(JOLNSIP Ч2эцодод) “JOA 39 `иЗО vINJIp ‘a “4 Хо} 94415-15910) “(MODSOY| JeOU) "1 sadjna ‘A *4 хоу мели э[ррии *(э^л1э$э44 уззоцолУ
‘eyyeyowey). ‘рр pupisuliag ‘A `Д ‘Хоу 1Арецу 148 03 Yo] мол 7] sadjna sadjn4 хоу pai JO 1010э ul иопемел э144е18098 Jo $94 А} BW “7 5384

7
ar)
|
го
1? т

Pa A
. au

ae
ently

es
ив

497

is all the more impossible to do. The reality of their differences
seems to be doubtful.

At the same time, the close relationship between our northern
foxes and the Scandinavian is quite evident and to this there are a
series of indications. Therefore, the form described here is provi-
sionally assigned to the nominal one.

2. Forest-steppe fox, V. v. diluta Ognev et Vorobiev, 1924.

Color lighter, dull and yellow, not so bright-reddish as in preced-
ing; fur shorter and coarser. Dimensions large—no less than in
northern form. Dimensions and proportions of the skull identical to
those of Moscow foxes (materials from Voronezh and Poltavsk
districts. V.E. Yakobi).

In forest-steppe zone of European part of the USSR.

Not recorded outside the USSR.

This form itself represents a transition from V. v. vulpes to the
following form.

3. Steppe fox, V. v. stepensis Brauner, 1914.

Color somewhat lighter than in V. v. diluta, fur shorter and
coarser. Measurements insignificantly less than in the preceding

form.

aa tue 1:7 )

. ИВ
И
Ne

gis | И ia eg
Ma al i Ly

Phot
ВАНО
: art

Fig. 122. Difference between tracks of Middle Russian fox V. v. vulpes L.
and Turkmeniyan И. у. flavescens Gray. Left—print of paw of Middle
Russian fox on shallow snow; track is broad and diffuse since pads are
densely covered with wool, claws quite short and blunt; vic. Moscow, Nov.
1945. Right—track of Turkmeniyan fox on wet sand, paw is smaller, slender,
nearly without wool below, claws long and fine; vic. Bakhardok well*,
Karakum, 28 Jan. 1948. Sketch by A.N. Formozov, about 2/3 nat. size.

*Misspelled in Russian original, “Bokhudok’—Sci. Ed.

338

498

In Pri-Chernomorsk and Pri-Azov steppes; in part in western
Cis-Caucasus steppes.

Not recorded outside the USSR; possibly encountered in Pri-
Chernomorsk [Black Sea region] steppe regions of Romania and
Bulgaria.

4. Crimean montane fox, V. v. crymea-montana Brauner, 1914
(syn. krymensis).

Similar in all respects to V. v. stepensis, but fur overall some-
what brighter, fluffier and denser.

In mountains of Crimea.

Not recorded outside the USSR.

The separation of this montane, chiefly montane-forest, form
cannot be considered well founded. Other characters, mentioned by
the describing author and accepted by Ognev (1931), are not well
established.

5. North Caucasian fox, V. v. caucasica Dinnik, 1914.

Color very variable, from reddish to red-gray and nearly gray,
such that in western part of range brighter foxes prevail, but in
middle and especially eastern part, grayer ones. Fur short and coarse.
Measurements are large, not inferior to those of above-described
steppe fox.

In mountains and foothills of northern Caucasus; in part on
adjacent plains, except Pri-Caspian part, and probably also south
slope of Main range, at least in west.

Not recorded outside the USSR.

Generally, this is well-defined race characterized by very great
mixing of characters and general amplitude of variation. On a par
with nearly typical gray foxes appearing in the east, bright foxes
with relatively good coats are met with in the west, in the Kuban’
basin. Most typical, apparently, are “red gray” foxes of the middle
part of both the ranges and foothills.

The above-mentioned particularity of this form is, it seems, to
a considerable extent connected with the fact that its features are
the result of mixing of characteristics of the true montane fox of
the Main Caucasus range with characters of foxes of southern
Russian and Azov-Black Sea steppes coming in from the west and
north, and with characteristics of karaganka (in the east) and of
montane Trans-Caucasian fox in the south. This form, apparently,
itself represents to a certain degree the result of interracial hybrid-
ization.

339

499

6. Eastern Trans-Caucasian fox, V. v. alpherakyi Satunin, 1905.

General color dirty rusty-gray or rusty-brown with brighter rusty
stripe along spine. Individual bright-reddish animals are encoun-
tered. Fur short, coarse, and sparse. Measurements small. Maximum
skull length of males 132—139 mm, females 121—126 mm; condylobasal
length in males 130—135 mm, zygomatic width in males 69—76 mm
weight to 4 Ко.

In plains of eastern Trans-Caucasus in Kura and Araks basins
on the west, apparently, to Tbilisi; valley of middle course of Araks;
Caspian coast north of Apsheron peninsula.

Outside the USSR—in adjacent parts of Iran and, perhaps,
Turkey.

This well characterized form belongs, according to characters
of size and color, to the group of the southern semidesert and
desert foxes—karaganka. Together with this, its features, in par-
ticular those of the skull, are not manifested to the extent as is
found in foxes of Turkmeniya living at the same latitude beyond the
Caspian Sea.

7. Trans-Caucasian montane fox, V. v. kurdistanica Satunin,
1905 (syn. alticola).

Color pale-yellow or light-gray, sometimes brownish-reddish.
Fur fluffy and dense—denser and fluffier than in other Caucasian
forms. Measurements large—larger than those of V. v. alpherakyi,
but not smaller, or only slightly less, than north Caucasian form,
Г. у. caucasica.

In Armenian highland and the other montane districts of the
Little Caucasus.

Outside the USSR, in parts of Asia Minor and Iran adjacent to
Trans-Caucasus.

A well-defined form.

8. Anadyr fox, V. v. beringiana Middendorf, 1875 (syn.
anadyrensis, kamtschadensis, kamtschatica, ? schrenkii, splen-
didissima).

Color saturated bright-reddish, almost lacking in bright ripple
along back and on body sides. Coat very fluffy and soft.

Measurements large, no less than in foxes of northern Euro-
pean part of the USSR.

33Available accurate data on dimensions completely inadequate.

500

In Anadyr territory, Chukotsk land, Kamchatka, Kuril islands
and Sakhalin. It is probable that only animals of the northern Kuril
islands belong to this form, and Sakhalin foxes are assignable to a
separate form (schrenkii), while those inhabiting the middle part of
the Kuril chain occupy an intermediate position between them (V.G.
Voronov and A.G. Voronov). This suggestion requires verification
based on good material.

Absent outside the USSR. It is possible that foxes of western
Alaska belong to this form (Middendorf, 1875).

The brightest form in the Old World.

9. Yakutsk fox, V. v. jacutensis Ognev, 1922 (syn. sibirica).

General color of back, neck and shoulders is brownish-rusty,
sides bright ocherous reddish-yellow. Coat very fluffy and silky.
Measurements large but, apparently, somewhat smaller than in
У. v. beringiana.

In Yakutia and Siberia southward to Baikal and Stanovoi range
and eastward to Okhotsk coast.

Absent outside the USSR.

One of the very brightest forms.

10. Trans-Baikal fox, V. v. daurica Ognev, 1931 (syn. us-
suriensis, dolichocrania).

Color along spine quite light, dull yellowish-reddish, with strongly
developed whitish ripple and grayish longitudinal stripes on anterior
side of limbs. Coat fluffy but somewhat coarse.

Measurements large, not less than those of Middle Russian fox.

In Trans-Baikaliya and Amur.

Outside the USSR—uin adjacent parts of Mongolian Republic
and, probably, northeast China.

Well characterized form, analogous in its geographical position
to that of European У. у. diluta and very similar to it in color. |

Nomenclature of V. v. daurica cannot be considered estab-
lished. If this form is distributed to Peking [Beijing], which is probable,
then it follows that it is named ¢tschiliensis Matschie, 1907. The
name huli Sow. given to Manchurian fox is hardly acceptable; it is
an evident /apsus for hoole—the name of an animal from Fukien.

The systematic position of foxes of Ussuri territory (Primor’e)
is not clear. It is not excluded that they belong to the described
form, although it is more likely that they differ. In that event, they
will receive the name dolichocrania Ogn., 1926 or kiyomassai
Kishida, 1929.

340

501

11. Tobol’sk fox, V. v. tobolica Ognev, 1926.

General color yellowish rust or dirty-reddish with well devel-
oped cross, and often black area on belly. Fur is particularly fluffy
and long. Dimensions large.

In plains districts of western Siberia in basin of middle and
lower course of Ob’ river.

Absent outside the USSR.

12. Karaganka, V. v. caragan Erxleben, 1777 (syn. melanotus).

Color light sandy-yellow or yellowish-gray. Fur short and quite
coarse. Measurements are somewhat smaller than those of Middle
Russian fox. In skull may be noted same above-described feature
as in “gray” foxes.

In steppe and semi-desert half of Kazakhstan, including Ustyurt
and Pri-Balkhash sands.

Outside the USSR, possibly in western Dzhungariya.

Forest-steppes of southern west Siberia and northern Kazakhstan
are, apparently, occupied by a separate form of fox representing a
transition from V. v. tobolica to V. v. caragan. This large fox is
quite light reddish-gray in tone; below a dark area is absent. Fur
somewhat rough, but quite fluffy. It is, apparently, related to V. v.
diluta.

In its turn the karaganka, features of which are quite variable,
itself represents a transition to the next form.

13. Turkmeniyan fox, V. v. flavescens Gray, 1843 (syn.
cinerascens, splendens).

Fox of “gray” color, very small size, with small infantile type
of skull (see above, Fig. 121 and Table 39). Smallest of all forms

Table 39. Skull measurements of Turkmeniyan fox (vic. Ashkhabad;

V.E. Yakobi)
Data Males Females
N. Min. Max. Aver. М. Min. Max. Aver.
Greatest length 14 129.3 145.5 134.2 121151282 143190134:
Condylobasal 14 1257 > 13911324: De D2 Rs 37:4: 1293
length

Length of upper 14 49.0 Me D2 12.1486 1246. 520
molar row
Zygomatic width 14 67.8 TOS AO ete 05 238 0691

Greatest width 14 44.7 49.6 46.2 12 440 477 46.2

502

inhabiting our country—and grayest, with maximum degree of re-
duction of reddish pigment. Body length 49-57.5 cm, length of tail
33.5—39.5 cm; ear length 7.7—10.0 cm; sole of hind foot 12—15 cm.
Weight 2.2—3.2 kg (Southern Tadzhikistan; Chernyshev, 1958).

In plains of Middle Asia approximately south of latitude of
Ustyurt and Aral Sea.

Outside the USSR—in contiguous parts of Iran and Afghani-
stan.

Within limits of USSR, one of most sharply distinguished race
of the species. Its relationship to forms described from the more
southern districts (see below) may not be. considered clear.

Ognev’s (1931) suggestion about occurrence of V. v. splendens,
described from Astrabad in northeastern Iran, in adjacent parts of
our country (in southwestern Turkmeniya) was not confirmed. Ap-
parently this form, described from one exceedingly bright specimen,
was generally not real. Among the typical “gray” V. v. flavescens
in southwestern Turkmentya, fairly bright ginger foxes are some-
times encountered (less than 1%) (V.G. Heptner).

14. Turkestan montane fox, И. v. ferganensis Ognev, 1926
(syn. pamirensis, ochroxantha).

Color quite bright light-reddish or light-yellow, rarely with gray.
Fur denser and fluffier (especially in high mountains) than in И. у.
caragan and especially V. v. flavescens. In size somewhat larger
than V. v. flavescens and similar to V. v. caragan.

In mountains of Middle Asia westward to the Amu-Dar’ya.

Outside the USSR—apparently in adjoining parts of Afghanistan
and China, and in part, apparently, in Tibet. It is most probable, that
the actual name of this form is montana Pearson, 1836**.

The systematic position of foxes from a series of territories still
remains completely unclear. Apparently, foxes of the Yenisei basin,
western Siberian forest-steppe and Altai are unique (Kuznetsov,
1944). V. v. schrenkii Kishida, described from Sakhalin is, appar-
ently, close to V. v. beringiana, or identical to it (fur standards for
the Sakhalin fox do not distinguish it). A distinct form, V. v.
splendidissima, was described for the northern and middle Kuril
islands. Information about these forms is completely insufficient
(see above, description of Anadyr fox).

“т the following notes, materials of Kuznetsov (1944) were used. In all
cases, it was the predominant color type of the species.

503

For foxes living outside the boundaries of the USSR, a great
number of names have been coined. Usually, the forms below are
recounted.

In Europe: 1) V. v. crucigera Bechst., 1789 (all Europe, except
Scandinavia, Spain and some islands of the Mediterranean Sea’; 2)
V. v. ischnusae Mill., 1907 (Sardinia and Corsica); 3) V. v. induta
Mill., 1907 (Cyprus); 4) V. v. silacea Mill., 1907 (Spain).

In Africa: 5) V. v. barbara Shaw, 1800 (Morocco); 6) V. v.
aegyptiaca Sonn., 1816 (Egypt, Libya); 7) V. v. atlantica Wagn.,
1841 (Algeria).

In Asia: 8) V. v. montana Pears., 1836 (Sikkim, Yunnan; Tibet,
Kumaon, Nepal, Punjab to Gilgit); 9) Г. у. pusilla Blyth, 1854
(Northwestern India from Punjab to Rajputana, Sind, Kutch,
Baluchistan, southern Iran, Iraq); 10) V. v. griffithi. Blyth, 1854
(Afghanistan, Waziristan, northern Punjab); 11) V. v. japonica
Gray, 1863 (Japan); 12) V. v. hoole, Sw., 1870 (southern China
from Sichuan to Fujian); 13) V. v. arabica Thos., 1902 (Arabian
peninsula and Syria); 14) V. v. tschiliensis Matschie, 1907 (north-
ern and northeastern (?) China); 15) И. у. апаюйса Thos., 1920
(Asia Minor); 16) V. v. palestinae Thos., 1920 (Palestine and
Lebanon); 17) V. v. peculiosa Kish., 1924 (Korean peninsula).

In America®; 18) И. v. alascensis Merr., 1900 (Alaska and
northern part of range eastward nearly to Hudson Bay); 19) V. v.
harrimani Merr., 1900 (Kodiak Island); 20) Г. у. kKenaiensis Merr.,
1900 (Kenai peninsula); 21) V. v. bangsi Merr., 1900 (Labrador);
22) V. v. deletrix Bangs, 1898 (Newfoundland); 23) V. v. rubricosa
Bangs, 1897 (region north of Great Lakes eastward to the ocean);
24) V. v. fulva Desm., 1820 (eastern and southeastern states east
of 100° W. Long. (see footnote 27 on page 482); 25) V. v. regalis
Merr., 1900 (western parts of Ontario, Manitoba, part of
Saskatchewan, and south to Nebraska and Iowa); 26) V. v.
abietorum Merr., 1900 (British Columbia, northern Alberta); 27) V.
v. macroura Baird., 1852 (region of the Rocky Mountains south of
British Columbia to New Mexico); 28) Г. v. cascadensis Mertr.,
1900 (southwestern British Columbia, Washington, Oregon); 29) V.
v. necator Merr., 1900 (Nevada, California).

Two “morphs” of this subspecies were described for Bulgaria—the moun-
tain (monticola) and plain (planicola) (Atanasov, 1958).

%Мапу American authors assign their red fox to a separate species, V. fulva
Desm., 1820.

343

504

Therefore, about 43—45 subspecies are considered within the
limits of the species. This number is, apparently, excessive. Leaving
aside forms examined by us, one may a priori note the doubtfulness
of the known number of forms of the Old World, and also perhaps
the American (though their number is not so great).

With all of that, the red fox remains one of the species most
remarkable as regards geographic variation (V.H.).

Biology

Population. In the Soviet Union, the fox is one of the most com-
mon predators. Proceeding from counts of individuals in restricted
areas and annual catch of 80% on the average (Chirkova, 1947 and
others), the general population in the territory of the USSR in the
post-war years may be determined approximately as several hun-
dred thousand to | million head and more (in summer after
reproduction in various years).

The number of foxes is not the same in various natural zones
and in different biotopes. The species is rare in forest zones with
prolonged snow cover and in regions of permafrost—in the north
and east of the European part of the Soviet Union, throughout the
taiga zone in Siberia and especially in the deep snow districts of
western Siberia and Pri-Yeniseisk territory. The fox is rare in tun-
dras—Bol’shezemel’sk tundra, in the southern half of Yamal,
Gyadansk and Taimyr peninsulas, and eastward. Some increase in
the population density of foxes in the forest and tundra zones is
recorded in the southern belt of shrub-tundra and in the forest-
tundra. The fox is comparatively numerous in some years in the
southern part of Novaya Zemlya, in Kanin, in the lower Kolyma, in
the middle and upper course of the Anadyr, in the mountainous
country between the Lena and Vitim, in the middle course of the
Vilyui, in the taiga of southern Trans-Baikaliya, in Kamchatka,
Sakhalin, in Ussurii territory, and on adjacent islands. There are
many foxes in the southwestern forest zone in the European part
of the USSR. The fox is especially numerous in the steppe zone,
in forest-steppe, in the deserts and foothills of the Caucasus, Trans-
Caucasus and Middle Asia, where snow cover is absent or does
not lie for long, the food is available throughout the year, and the
reproductive conditions are favorable (Fig. 123).

‘(RAOYMYD “A'W) $148 001 0} $19}21 10q ‘(леэА Jad э8влэле) 66/5661
—CZ/p76| WoO зречэуеш [е91$14е$ “vole lun Jod sulys Jo р|э1А uo poseq YSSN Jo Алоэ} ио иоцетдо4 хо; Jo Aylsuaq “¢€Z] ‘914 TPE

051 оп

у ae 08
А
ШУ 009 — 006

с
ИТС

— SS SS
ee

2
==>

| >
м

и ae
iil

ice
=a
———
———_ .
=—
=
А

a =
SS -
SES

344

506

The density of fox populations in the pre-tundra and forest
zones is characterized by the following indicators. In Kanin [pen-
insula], in the neighborhood of Lake Propashchi, only one fox was
recorded when drive-hunting in an area of about 400 km? in a year
with minimum numbers (0.03 fox per 100 hectares). On a 10-km
route in Pechoro-Ilychsk preserve, for a series of years, only 0.07—
0.6* fox tracks were encountered; in the Kharovsk region of
Vologodsk district—3.3 tracks on the average, fluctuating in differ-
ent years from 1.4 to 6.2 tracks (1944—1950); on the Zavidovsk
farm in Kalinin district—1.0—7.8 tracks, in Kirov district—0.5—7.0
tracks, but in summer after the appearance of the young 2—3 foxes
on 1,000 hectares. In the Pogonno-Losino-ostrovskoe forestry near
Moscow, there were 5.8—28.3 tracks on a route of 10 km (average
of 20 years), and on 1,000 hectares, an average of 2.1 foxes were
counted in winter. In the Smolensk district, in 1,000 hectares, 4 and,
in some places up to 10.5 foxes, lived (Table 40) (Teplov, 1949;
Chirkova, 1952; B.A. Larin, B.A. Pestov and others).

In the Pri-Yeniseisk taiga, in the middle of the previous century,
on a 10 km. route, an average of 0.02 to 0.12 fox tracks were
encountered in various years (Middendorf, 1853). In the Magadan
district in 1956—1959, on 1,000 hectares of various areas, 0.08—2.0
foxes occurred (Shustov and Belozorov, 1959). On a 10 km route
in the Irkutsk district and in the Buryat ASSR, 0.3—3.0 tracks were
counted, while in Karagin region of Kamchatka district—0.2—2.3
tracks; in the latter, on 1,000 hectares, 0.2—0.8 and more foxes
were met with. In Ul’chsk region of Nizhneamur district, on a 10-
km route, 0.2 tracks were encountered and on 1,000 hectares, 1.8
foxes. In the Amur district, on a 10 km route—0.5—3.6 tracks, in
Primor’e territory—2.0 tracks, and оп 1,000 hectares—1 fox.

One fox burrow was found in an area of 290 hectares near
Moscow in a forestry plot of 2,000 to 3,000 hectares in southern
Vologodsk district and in the forests of the Far East.

In the steppe zone and semideserts in the southeastern Euro-
pean territory of the Soviet Union, the population density of foxes
increases 10 times, compared to the taiga region. The fox is espe-
cially numerous in the broken steppe landscapes of the Cis-Caucasus,
in the Crimea, in the southern Ukraine, in the eastern Carpathians,

*Probably refers to per 10 km—Sci. Ed.

344

507

Table 40. Index counts of fox populations
Foxes Tracks in a Burrows
Zones, districts in 1000 in a 10-km !0-km route Е ae
hectares route те
Forest 0.2—3.0 0.12.0 0.02—7.8 0.3—4
(to 10.5)37 (to 28)38
Forest-steppe and 0.9-33.0 0-10 2.556 1-25
steppe (and to 48)°9 (and to 83)”
Semidesert and desert to 24 0-3.5 0-20 2-11
Montane regions of 2-20 0.52.0 0.5—37.5 12—25
the south (and more)*?

on the right bank of the Don, in the central chernozem belt, in the
Orenburg steppe, in the forest-steppe and steppe of the Altai,
Minusinsk and Tuvinsk depressions, and also in southern and east-
ern Trans-Baikaliya, in the steppes of western Siberia and
Kazakhstan, and in the semideserts and in part the deserts of Middle
Asia.

In the steppes of the European territory of the Soviet Union,
fox numbers are greater than in the Asiatic part of the USSR (Izmailov,
1940; Chirkova, 1941, 1947a, 1952a; V. Popov, 1953; Malenkov, 1953;
Obtemperanskii, 1955; I. Romanov, 1956; М.М. Bakeev). In a 10-km
route, in the steppes of Stavropol’sk territory (at the end of the 30’s),
an average of 44—56 tracks were encountered, in the Usmansk pine
forest of the Voronezh district (in 1948—1953) —from 1.0 to 21.8 tracks.
In 1000 ha, in summer, together with the cubs in the litters, foxes
amounted to: 0.9 to 3.8 in Brovarsk region of Kiev district (1947—1952);
in Verkhnekhavsk region at the edge of Voronezh preserve—3—16 (1949—
1950); in Staromar’evsk region of Stavropol’sk territory in summer
from 3.5 to 12.5 and in winter, 10.3—16.5 (1938—1941). In separate
biotopes of the latter region, up to 3—5 foxes were counted in 100* ha,
while in the beginning and middle of the 40’s, in one place, in winter,
around 17—20 and even 30 foxes might be observed, mousing in packs.
In the absence of a harvest in the course of a series of years in

37In Smolensk district.

38In Podmoskovsk Forestry plot.

In small areas in Stavropol’sk territory.

“In small parts of former Taldy-Kurgansk district.
*Should be 1000 hectares?—Sci. Ed.

345

508

Arzgirsk region of Stavropol’sk territory, on 1000 ha, from 8.8 to 26
foxes were counted in summer of 1938—1941. At the end of the 30’s,
in Khopersk preserve from 2.5 to 3.5 litters were distributed on
1000 ha in summer, and in winter—10 foxes. In the steppe of
Uzhursk region in Krasnoyarsk territory, 2.4 foxes occurred on
1,000 ha.

In desert and semidesert zones, foxes are numerous in many
places. They are especially many on the Priergeninsk plain, along
the Volga-Ural interfluve, on Ustyurt Plateau, in the Sarysu valley,
in southern Pri-Balkhash, and also among the sandy deserts of
western and southern Turkmeniya—in Balkhan [mountains] and
throughout the entire lowland section southward to the lower Atrek,
in Maryisk and Kerkinsk districts, and in the foothills of Kopet-Dag
in southern Turkmeniya. Foxes are relatively scarce in Betpak-
Dala, in Muyunkum and along the Syr-Dar’ ya.

In the Kzyl-Ordinsk district, along a 10-km route an average of
4.1 fox tracks were found (winter of 1940/41); in the northern
Kyzylkum and on the coast of the Aral Sea, 0.3 foxes were counted;
in southern Ustyurt—0.3 and in the northwest—1.3 animals; in the
lower Amu-Dar’ya, only 1 fox was met. with in spring, and in
autumn, 1.5 foxes; in former Bel’agachsk region of Semipalatinsk
district, in March 1943—3.5 foxes; along the middle and lower course
of the Murgab in March, there were more than 10 fresh tracks and
a carcass. In the Betpak-Dala desert, in the course of several days
not even one fox was encountered. In southern Turkmeniya
(Badkhyz), in autumn 1935, 10 foxes might be taken from a ravine
near a spring. In Badkhyz preserve, in 1950—1958 (in an annual
route of 786 to 4,392 km), in each 100 km of the route, from 1.6
to 7.1 foxes were met with in spring, and in winter from 3.6 to 6.9
head. In Takhta-Bazar region, in 1943, during the day in winter, up
to 5 foxes could be seen within shooting distance along an 8-km
route. On the Ural-Emba interfluve, in an area of 400 km? an
average of 0.3 red and corsac fox litters were encountered per
1000 ha. In Balkhashsk region, in 1,000 ha, there were 0.5 litters;
in the vicinity of the city of Gur’ev—2.7 litters (23.2 foxes); in
Badkhyz preserve, up to 4 litters (24 foxes) (Kashkarov, 1935;
Kolosov, 1935; Rozanov, 1935; Heptner, 1956; Kostin, 1956;
Shcherbina, 1961; I.N. Nesterov, А.Е. Chirkova).

Foxes are numerous in the mountains and southeastern foothills
of the Caucasus and in the Trans-Caucasus, in the Alai mountains

509

and in the western Pamir, in the Karatau range and along the
Pyandzh, in the high-montane belts of Tien Shan (on “syrt”* of the
Tien Shan at an altitude of 3500 m above sea level, the fox is
common, and in places, even numerous; N. Naumov), in the
Dzhungarsk Alatau, in the southeastern Altai, especially in its south-
western foothills.

In the subtropics of the Black Sea coast, foxes are rare, as well
as on the southwestern slopes of the Caucasus, in the eastern
Pamir and in the forest zone of the Altai and Sayans. Fox is absent
in the nival zone and, apparently in the “cold desert” of the Tien
Shan.

In the Saraibulagsk, Aiotsdzorsk and Uritsk ranges in Armenia,
from 2 to 20 foxes were recorded in the spring and summer in a
1000-ha territory. In the Pambaksk range and in the vicinity of the
city of Erevan, in January—March, an average of 3.7—6.0 tracks
were met with along a 10 km route; in the vicinity of the city of
Kirovakan—36 tracks; in the mountains of the Kirghiz range—37.5
tracks. In the foothill regions of the Altai, in summer of 1946, a
year with relatively abundant foxes, 5 litters were found in a ter-
ritory of 5 km? (Dal’, 1944, 1948, 1950, 1954; Chirkova, 1952a;
V.N. Belyaev, M.A. Kuz’mina).

Throughout the historical period, fox populations have changed,
mainly under the influence of human activity. Thus, in Kamchatka
in the 17th century, Krasheninnikov (1755) found such an abun-
dance of foxes that it was sometimes necessary to beat them back
with a stick from troughs where the dogs were fed. Many foxes
were reported on the Crimean peninsula by Gablitsl’ (1785). A.M.
Nikol’skii (1889) spoke of the great quantity of foxes on Sakhalin
during the past century. [Human] population growth on Kamchatka,
Sakhalin and other regions of the Far East and extreme North
caused fox harvesting to develop and to a decrease in their num-
bers. In the deserts of southern Turkmeniya (interfluve of Tedzhen
and Murgab), where the number of foxes is now very high, their
number was higher—in the winter of 1886/87, a group of hunters
caught 14 thousand foxes and in the preceding year 12 thousand
(Heptner, 1956). But, in the greater part of the territory of the
Soviet Union, growth of population, expansion of areas under cul-
tivation, thinning of forests in the north, sowing of tall-stemmed

*Elevated watershed.

346

510

plants and forest-plantations in the steppe zone, and irrigation and
acclimatization of deserts helped to increase fox populations.

Zoologists of the past century frequently noted the low number
of foxes in the forest-steppe and steppes of western Siberia. From
the mid-’40s of the 20th century, the number of foxes began to
increase. Tanning of fox skins in Chelyabinsk, Kurgansk, Omsk,
and Novosibirsk districts, and in the Altai territory at the end of the
first decade and in the 50’s increased several times.

Habitat. Red foxes everywhere prefer open and semi-open
expanses. In the tundra and forest-tundra zones, foxes usually are
restricted to the edges of riparian woods, among bushes of floodlands
and valleys, along lake shores and rarely /aida on the seashore. In
all seasons of the year, it goes out into the open tundra zone; it is
regularly encountered in the subzone of shrub and cotton-grass
tundra, preferring elevated relief. In the moss-lichen subzone of the
tundra, they appear sporadically. Among the Arctic tundras, the fox
is common only in the southern regions of Novaya Zemlya and in
the Far East. In connection with climatic warming in the Arctic in
the course of the last decade, movement of foxes into the tundra
has been noticed (Skrobov, 1958, 1960a). In Magadan district, the
fox serves as the main species exploited in the high-bush tundra
and is the second species in the zone of Arctic tundra. The distri-
bution of foxes in 1000 ha plots (in a southwestern direction)* in
Magadan district is as follows (Shustov and Belozorov, 1959):

Tundras, bushes and swamps of the central

subzone of the Arctic tundras 0.08 foxes
Tundras, bushes and swamps of the southern

subzone of the Arctic tundras 0.13 foxes
Inundated broad-leaf forests and bushes 1.0 foxes
Inundated mixed forests 2.0 foxes

In the taiga zone, the fox is rare in the depth of the great,
remote plain of tall-trunked forests where large areas are occupied
by mossy bogs, where soft snow lies for a long time and is very
deep. The fox prefers diverse, mixed-age forest among broken
relief, and alternating with fields, meadows, grassy swamps and

*Appears to refer to the list of vegetation types, from northeast to south-
west—Sci. Ed.

Sila

river valleys. It selects biotopes where open areas prevail in au-
tumn and winter. In spring and summer, in the breeding period
foxes are concentrated in more remote forest sections. In winter,
in Karelia, tracks were encountered along a 10-km route (Marvin,
1959) [as follows]:

in pine forests with berries —

in pure pine forests 0.43
in pine forests with sphagnum 133
in mixed pine and spruce forests —
in sedge swamps 7.4
in clearings and open burns 7.84
in broad-leafed forests 14.8
average index of occurrence 2.05

In Kharovsk region of Vologodsk district, at the beginning and
end of winter, on routes of more than 500 km, 14.7 fox tracks were
encountered per 10 km in clearings, 12.4 in sedge swamps, 9.8 in
young coniferous forest. From 6.6 to 4.7 tracks were counted in
pine forest in a sphagnum swamp, in fields, open swamp, young
mixed forest and in meadows; in a young broad-leafed and a mixed
grassy forest, 2.8 tracks. The lowest number of foxes visited dense
firs—1.2 tracks per 10 km route (B.A. Larin).

In forests in the central zone of the European part of the

- USSR, foxes hold to various biotopes. From the Byeloruss and

347

Novgorod districts in the west to the Udmurt and Kirov districts in
the northeast, the fox is restricted to 72% of cases in autumn—
winter time to meadows, fields along edges of forests, beside open
swamps among clearings, and floodlands of rivers and creeks. Foxes
are encountered in approximately the same biotopes in winter in
Sverdlovsk and Chita districts and in the Buryat ASSR.

In the forest-steppe and steppe zone, the fox may be met with
in all biotopes. It often inhabits places rich in hills and ravines, with
access to rocky aspects or with bairach* forests and also steppes
and fields of various kinds, intermixed with islands of forest or
thickets of shrubs. On flat steppes in summer, the fox makes wide
use, for breeding and shelter from danger, of all unevenness
of fields and sown tall-stemmed cereals and technical culture—

*Bairach refer to forests in deep, flat valleys within the steppes—Sci. Ed.

sunflower, maize, castor bean, wheat, and winter rye, as well as
thickets of weeds, ground cherries, isolated groves of birch, aspen,
willow, and forest plantations of the broad-leaved variety. After the
gathering of the harvest from the fields, the fox hunts even in
entirely open biotopes and rests and hides from danger in beds
overgrown with weeds, in reed beds, in forest belts, old pine
forests, etc.

In the southern zone of grass-forb steppes of Stavropol’, aver-
age density of foxes in 100 ha (end of the 40’s) was in stubble, 5.8
foxes (from 0.3 to 24.0), in fields of sunflowers, castor bean and
maize—3.7 foxes in summer and considerably more in autumn; in
weeds—3.5 (from 1.3 to 5.6), in speargrass-sagebrush steppes with
open gulleys and rocky hillocks with osier—3.3 (from 0.4 to 16.7),
in broad-leaved forest, in greenery and in forest gorges on average,
1.2—1.8 (from 0.1 to 60.7; in forest gorges, in some years in autumn,
6—7 foxes were bedded per 10 hectares; М.М. Bakeev).

In the desert and semidesert of the Pri-Caspian, in southern
and western Turkmeniya, the fox prefers sandy biotopes with a
network of sandy ridges. It is met with particularly often among
undulating sand-solonetz of lowlying steppes, grass-sagebrush deserts
and steppes, saltsage-sagebrush deserts, and wormwood-turf steppes,
in Ustyurt and in the Balkhan [mountains] along precipices and
gorges of the low steppe mountains with thickets of various bushes,
in saxau! forests and in riparian tugais [woodland] along Middle
Asian rivers interspersed with open expanses of sands and of
dzhingiliya [Halimodendron halodendron], Russian thistle, tama-
risk, pea-shrub [caragana] and other shrubs. Traces of active
movement of foxes can be observed among sand dunes and in hills
covered in spring with rich grassy cover on the lower course of the
Murgab, and also in the impenetrable thickets of reeds and in pis-
tachio groves in the rocky passages in southern Turkmeniya.

In Badkhyz preserve, the fox is also unevenly distributed. For
1950—1956, along a general route of 55,992 km, foxes were encoun-
tered per 100 km as follows (Shcherbina, 1961):

in desert-steppe biotopes 1.2
in the Kushka river valley 5.8
in pistachio groves 6.5

average for 2,120 foxes 3.7

348

513

Foxes become seasonally redistributed among biotopes. In win-
ter, food attracts many animals and in summer, waterholes on the
Kushka river (Heptner, 1956).

The fox is rare in clayey sagebrush plains of semidesert and
deserts, and in waterless solonetz depressions of Betpak-Dala.
According to counts in the winter of 1940, in Kzyl-Ordinsk district,
along a route of 1 km with a light snow fallen 4 days before, 3
tracks were found among the scarce saxauls, in an overgrown
clearing in saxaul—1.2, in clearings in saxaul and solonetz steppe—
1; in 3 km through old saxaul clearings in hilly desert, and along 50
km in reed thickets, not one track was found (B.M. Kasatkin).

In the Caucasus, the fox inhabits valleys, and mountains to an
altitude of 2500—2800 т; it lives in the foothill-steppe, montane
forest zones, and montane meadows of the subalpine and alpine
zones. It is found in low growing forest, shrubs, and reeds, in
weedy areas, fields and open steppes; it prefers rocky and stony
places. The fox does not go into the nival zone above 2800 m above
sea level. Foxes are rare in the forests of the foothills of the Black
Sea coasts and in the tall-trunked beech forests of the southern
slope of the Great Caucasus. In Azerbaidzhan, foxes were encoun-
tered in virgin semidesert in 59% of cases; in low forests—25%, in
beech-hornbeam forests on the southern slopes of the Caucasus—
in 6.8%, and in alpine meadows in 3.1% of the cases (Vereshchagin,
1952). In the Pamir-Altai and Tien Shan mountain systems, the fox
inhabits all vertical zones, but populations are especially dense in
subalpine and alpine meadows to an altitude of 3000—4000 т, ap-
proaching the limit of permanent snow. In the Altai and Sayans, the
fox is, in the majority of regions, rare, and is restricted mainly to
steppe and forest-steppe sections.

In years with food shortages, foxes approach human domiciles.

Food. The fox is an omnivorous predator. In the Soviet Union,
up to 300 animal species and several tens of plant species on which
it feeds are known. Small rodents constitute the main food of foxes.
It feeds most often on various species of gray voles of the genus
Microtus; in their absence, other species abundant in a given region
predominate in the food. Small rodents constitute in the fox diet
anywhere from a single individual to 100% of dry weight, and rarely
less than several tens of a percent in an analysis. The fox feeds
less on birds, of which passeriforms, galliformes and waterfowl

514

predominate. It often feeds on insects and other invertebrates, as
well as on plant food—fruits, berries, seeds and vegetative parts of
plants, and along the banks of river and inland waters—on fishes,
and along the coast—flotsam (marine animals, fishes, echinoderms
and others).

_- Their food varies in different geographic landscapes (Fig. 124).
In the pre-tundra belt and taiga zone, the northern voles Microtus
middendorffi, M. oeconomus and M. agrestis serve as the chief
food, and rarely, species of the genus Clethrionomys. Mouse-like
rodents were encountered in food analysis as follows: up to 93% in
Lapland preserve, to 70% on Kola peninsula, 62% in the Middle
Kolyma region and Kamchatka, 90% (of 100% occurrence) in eastern
Sayan (“Stolby” preserve), 35—38%, during an abundance of small
rodents in the Middle Sakhalin preserve (Nasimovich, 1948; Dul’keit
and Kozlov, 1958; L.P. Borodin, A.I. Zubov, A.V. Kondratov, A.N.
Shcherbakov, E.P. Volkova). In the extreme North the fox, in the
absence of gray voles, destroys a great quantity of lemmings of the
genus Lemmus (up to 67% in winter in Lapland preserve).

With a scarcity of small rodents, it catches forest game birds
(up to 40% occurrence in Lapland, 24.6%—in Pechoro-Ilych, 12—
14% in Middle Sakhalin, and 5.3% in Sayan (“Stolby’’) preserves);
willow ptarmigan, capercaille, hazel grouse and black grouse. In
same years, it often catches white hares (up to 15% occurrence in
Lapland and up to 52.7%—in the Pechoro-Ilych preserves). The
forest game birds and white hares are consumed in great quantities
only in years of epizootics and pestilence among them. Especially
many white hares were devoured by foxes in Yakutiya in years of
massive epizootics and die-off of this rodent* after its legendary
reproduction in 1955—1957. At that time more than half of the foxes
examined had fed on hares (more than 50% occurrence in analyzed
food and 96% occurrence in feces of 2 fox litters; Labutin, 1960).
In other years, foxes consume hares comparatively rarely (in Middle
Kolyma region 3.9% occurrence in food, in Middle Sakhalin pre-
serve in summer, 8% and in winter 6%; in Kamchatka and in
‘“Stolby” preserve, in analysis of some tens of stomachs and feces
of foxes, hares were not encountered, and only in preserve territory
were found in 2 cases out of 8 remains of fox feeding).

In the Far East, the massive migrations of salmonids—chum,
humpback, silver and others, which are stranded on river banks at

* Sic—Sci. Ed.

349

515

the time of spawning are of significant importance for foxes. Com-
position of the food of foxes on the Kuril islands is unique; on Urup
Island, in August 1955, the main feeding areas were along the
seashore. In feces and stomach contents there was no predominant
food; it consisted mainly of flotsam and the only rodent there—the
gray [Norway] rat— and also insects (in 24 fecal and stomach
content samples, % occurrence was as follows; Shmeleva, 1958):

gray rat TS
birds 88
fishes 65
insects 100
crustaceans 72
sea urchins 57
plants 83

Of constant occurrence in the food of northern foxes are sedges,
grasses, needles of spruce and fir, hazel [Corylus] and Japanese
stone pine [Pinus pumila] fruits. Foxes consume great quantities of
berries of crowberry, red bilberry, huckleberry, blueberry, rasp-
berry, mountain ash, rose, hawthorn and others.

In the southern belt of the forest zone of the European territory
of the Soviet Union, mouse-like rodents, in years when they are
scarce, are not always used as the main food. Besides all of the
animals possibly inhabiting the forest, foxes feed on carrion (up to
50% by weight and up to 100% occurrence in food analyses in
regions near Moscow in some years), usually around human dwell-
ings, in poultry yards, on garbage, eating edibles, and even unedible
wastes—scraps of leather, rags etc.

In the forest-steppe and steppe zone, the fox feeds almost
exclusively on small rodents, the chief of which consist of ground-
dwelling individuals which remain active throughout the entire year.
In the Ukraine, mouse-like rodents are encountered ш 91.3% of fox
foods investigated; of nine species, that which was eaten more
often than others, within a 7-year period, was the common vole
(Microtus arvalis)—75% in winter and 62% during the whole year
(Korneev, 1956; see below, Table 42). In the Stavropol’ steppe,
during a 4-year period, only 2 vole species were met with as food
common and social (М. socialis) voles—in 95—100% of cases in
various years (see below, Table 41). In the southeastern European

516

territory and in the steppes of western Siberia and Kazakhstan, the
particular species that serve as food are the narrow-skulled vole
(Microtus gregalis) and the sagebrush vole (Lagurus lagurus); in
Tselinn territory (former Akmolinsk district) both species in June
1946 constituted 90% of occurrences in the food (A.F. Chirkova).
In the forest-steppe of Novosibirsk and Omsk districts, mouse-like
rodents, during a period of sharp change in their number, consti-
tuted from 17 to 84% of occurrences in the food of some tens of
foxes by season (L.A. Popova, A.F. Chirkova).

In warm times of year, the fox hunts various species of the
gray ground squirrel [Spermophilus sp.]. Their remains were en-
countered in the food of the fox as follows: in the Ukraine, 2.3%
(Korneev, 1956), in the forest-steppe of Voronezh district—up to
38%, in the black soil forb-grass steppe of Stavropol’ territory—
13%, in the northern forb-grass steppe of Saratov district—47% and
in the semidesert virgin sheeps-fescue-needle-grass steppe of
Arzgirsk region in Stavropol’—67% (А.Е. Chirkova). In forest-steppe
regions of Novosibirsk and Omsk districts, the fox excavates ground
squirrels even in winter (8% of occurrences in stomachs; L.A.
Popova).

_ Snakes and lizards were found in the food of foxes in 30% of
occurrences in the Cis-Caucasian steppes. Insects (mainly beetles
and grasshoppers)—in Stavropol’ territory in a series of years—30%,
and in separate seasons of the year in Stavropol’ up to 62%, in
Kiev district—60%, in Saratov district—30%, and in Tselinn territory
(former Akmolinsk district)—50%. Of the plant foods, the fox eats
fruits of several fruit-trees and cultured melons, and in years of
rodent scarcity, it avidly eats rose hips which are rich in vitamins
as well as the wheat grains and sunflower seeds (V. Popov, 1943;
A.F. Chirkova).

The various species of gerbils constitute the main food of desert
foxes, depending upon their natural abundance. In the Volga-Ural
sands, foxes fed mainly on midday gerbil (Pallasiomys [=Менопе$]
meridianus) and tamarisk gerbil (Meriones tamariscinus). In the
Kyzylkum and Karakum, foxes usually hunt the great gerbil
(Rhombomys opimus); however, in spring 1946, in the sands of the
lower and middle course of the Murgab river, they fed more on
midday gerbil (61% of occurrences in feces) and more rarely on
great gerbil (25%). Dry weight of remains of gerbils in food con-
stituted 84% (A.F. Chirkova). In the lower course of the Ili river

517

in spring and summer of 1939—1941, half of the foxes examined fed
on four gerbil species, most often on great gerbil—5S—40% in various
years, more rarely on tamarisk gerbil—up to 17%, then midday—aup
to 10%, and red-tailed—up to 15% (I.G. Nitsenko, V.N. Belyaev,
A.A. Sludskii). In southern Turkmeniya, in years of Afghan vole
(M. afghanus) reproduction, the fox feeds mainly on it (up to 80%
of occurrences in food in the territory of Badkhyz preserve in
foothills of Kopet-Dag; Shcherbina, 1958). Among jerboas, foxes
more often hunt the thick-tailed three-toed (Scirtopoda telum) and
the large species of the genus Allactaga. Tolai hare [Lepus tolai],
which is usually difficult to find in the southern Pri-Balkhash, is
often hunted by foxes in years of its epizootics (48% of occur-
rences in feces in June 1939; A.A. Sludski1).

In the years when small rodents are scarce, the main substitute
food (though not as fully valuable) is served by insects*'. In north-
ern semideserts, they constitute in some years up to 70% of
occurrences in food analyses. In the deserts of Kzyl-Ordinsk dis-
trict, the fox feeds, in addition, on solifugids, scorpions and tarantulas
(up to 45% of occurrences in food around Lake Telikul’); in south-
erm Turkmeniya, foxes feed heavily on termites, beetles, grasshoppers
and others—up to 97% of occurrences of all invertebrates in food
of foxes in pistachios (1956) and up to 98% in the Kushka river
valley (1954); aggregate for 5-year period—in 70.8% of cases. In
Uzbekistan, insects were met with in food on an average of 70.9%
over 5 years. The red fox fairly frequently feeds on lizards, turtles,
snakes and their eggs, destroying many vipers in years of their
mass appearance. Plant feeding is also of great importance, con-
stituting in pistachios of Turkmeniya an average of 12.8%, and in
some years up to 48% of occurrences in analyses. Chief among
these is the pistachio nut—up to 74.8% of occurrences, and also
capers, watermelon, various grasses and seeds. In Uzbekistan, the
plant food constituted over 5 years an average of 74.5% of occur-
rences in the food, of which the main ones are: grape (20.2%),
grass, wild olive and others (Shcherbina, 1958, 1962; Ishunin, 1961,
1963; Formozov and Osmolovskaya, 1963). In the Tersko-Kumsk
sands, in October 1957, fox ate an abundance of ephedra fruits
(Ephedra sp.; Bakeev, 1959).

“With such food, reproduction of foxes falls (Chirkova, 1948).

350

Fig. 124. Components of food
of foxes in various natural zones

N
—
со

of Kola

stomachs, intestines, feces, food
tundra
Peninsula (Lapland preserve;

remains near the burrows, etc.).
I—Lightly wooded forest and

and seasons of year (contents of

montane

НГ; В; winter

|; В; winter

; winter

т
О

dra of former Srednekolymsk

forest-tundra and montane tun-
territory

Nasimovich, 1948; II—Deciduous

Zubov);

(А.

П-—Момапе taiga of eastern
Sayan (“Stolby” preserve near

оз
ая! ©
ava
ina} n
Sos
Ae
> —=
‚© | 2 =
N > o
om as
м о
AS
со > ©
m За
м9 S
(7)
We o
3 .-U 6
ar oe
оо 8
я м GS}
N« зо
оО Fa
о
мазя

V; В; summer-winter

5
=
=
&
&

spring-summer-autumn

.
9

@uUO0Z }$9.0-1

central and northern districts of
the European part of USSR
(A.F. Chirkova); V—Coniferous

forests (taiga) and Pacific Ocean

sheeps-fescue-needle-grass

Os > omy
Зоо
[96]
зови
Е ее
ОЕ =
Fok ad o
ga Cs?
ey So
lay | AS . Е З
о она
Saw = NS
eG SoS
Oo Te
ао ©
Oe & oe
o at 2 | SD
= О
= | gay AD) te)
ана
>>

steppe of Cis-Caucasus

"рэчииехо $}ээГ4о JO JaquNnu 94}
03 злэрэл WRISVIP YOR JO эцизэ
UI эл ‘sjewWuUe sUIeU—+y]
‘зитешэл э[91трэш рие рэцпиэрши
—¢] $348 ЧТ ‘soye1qoyJOAUl
JOyJO—| | :5$12э5 4+ —0]| :594$4-—6
‘sueiqiydue—g :5914э-—/ ‘зриа
—Q еше SNOJOATID9SUI—¢
‘SOIOAIUIVD §=[[eWIsS —p ‘UuOoll
-169—6 ‘soiey— ‘$19498—мэзэр
ит {sajoA Апмеш ‘5зиэро—|

$01119
теэи зитешэл э|адтрэит JO %—)
‘S9DUILINIIO JO % ИЕ Я ‘suleWal
ay JO з481эм Азр 40 % и у
(елодичо “AV ‘ешэцыу Аэ|ел
154242!) $58918 )--$И64 |, sue)
-цош YSIH—X] *(влозицо “A"V)
[1элы] де3апи| 19м9|-—94 ‘(7961
‘ви191эЧ94Ч$) э^195э44 zAYy peg
—e сАшэшуаТ (1910$
30 51155э9А- ША “(влохичо
"У ‘А10иа4э} 10904^81$)

Jee JO suosees |8 М ИЛ

о
и

Buds ‘у ‘ал

sowiwns-Huuds :5 ИЛ

Jouuuns-Buyds ‘У ПА

eS
KS

unos о surejunow рие syeseg

sedde)s pue sedde}s-jses04

Ise

353

520

In the mountains of the Crimea and Caucasus, the fox also
feeds most often on small rodents. In the mountains of Armenia
near Kirovakan, they constituted from 50 to 98% of the dry weight
of remains in feces and from 65 to 100% of occurrences in various
months from 1936—1939. They ate gray voles of the genus Microtus
more than others, mainly М. arvalis, but also quite often М. socialis
which, together with woodland voles (genus Pitymys) and the lo-
cally rare snow vole (genus Chionomys), constituted 85% of
specimens (of 1727) of all mammals in these investigations. Cases
of feeding on brown hares were rare (less than 0.1% of dry weight
of remains and 1—1.5% of occurrences). In all seasons of the year,
although not many times, fox hunted insects (up to 42% of occur-
rences), mainly locust, beetles, and grasshoppers (A.F. Chirkova).
In the Crimean mountains, the fox attacks young and weakened
individuals of adult roe deer (up to 29% of occurrences in food;
Kotovshchikova, 1936). In orchards, they select windfalls of apples,
pears, cherry plum, bird-cherry, blackthorn, grape and others; they
eat musk- and other cultivated melons.

Catching small rodents becomes very difficult with snow cover,
especially in the second half of winter with much snow, and also
after a thaw, when an ice crust forms in the course of a winter of
intermittent snow cover. The fox usually cannot break these crusts,
and because of this catching of rodents is delayed. In such years,
foxes turn to any living food available to them, and in their absence,
to carrion, refuse from human households, and even to feces of
animals and humans. Everywhere, in years of starvation, the fox
consumes small insectivorous mammals—shrews, water shrews
[Neomys]| and moles—which are most frequently only caught and
discarded, and usually not utilized by it as food, and sometimes
(even more rarely) feeds on small carnivores—weasels, ermine,
even marten and others, which usually constitute about 1% of oc-
currences among its food remains. However, in the mid 30’s, when
numbers of small rodents were low in central forest districts of the
European territory of the Soviet Union, small carnivores were eaten
by fox often, and, for instance, were found in the stomachs of four
foxes (of 12 caught). In these years, the stomachs of foxes were
at times completely full of hedgehog remains, swallowed along with
the skin as well as quills of these animals, which protruded into all
sides, bulging out from beneath the thin wall of the stomach (A.F.
Chirkova).

521

Age differences in foods of foxes are insignificant. In the
Stavropol’ steppes, and in the deserts of southern Pri-Balkhash,
food of young is more variable than adults.

In foreign countries foods are also varied. In Bulgaria, small
mouse-like rodents serve as principal foods, and more rarely, hares
(11% of occurrences in diet, part of which consisted of cripples),
then carrion, less—various birds; in summer—insects and other food,
quite often—fruits, berries, cultivated melons (Atanasov, 1958). In
Finland, the fox feeds mainly on domestic garbage and small mam-
mals (hares 6%, other species—less; Lampio, 1952). In England,
they often feed on rabbits, rarely on hares, and also on refuse and
carrion; in the absence of rabbits, mouse-like rodents and carrion
appear as the main foods (Lever, Armour and Thompson, 1957). In
the eastern and middle states of the U.S.A., the food of the red fox
consists of rodents, insects, birds, carrion and several other animal
groups. The main food is mouse-like rodents of the genera Microtus
and Peromyscus. In separate regions, muskrat, pheasants, grass-
hoppers and fruits are significant. In forests of the state of Michigan
(Murie, 1936), the fox often eats hares* of the genus Sylvilagus
and voles of the genera Synaptomys and Microtus, as well as
beetles, etc. (Nelson, 1933, Hamilton, 1935, 1943, 1944; Errington,
1935, 1937; Findley, 1956 and others).

The food composition of fox greatly varies in one and the same
place by year and season, and also by biotope, depending upon the
abundance and availability of one or another food. In the cher-
nozem [black soil] herb-grass steppe of the Staromar’evsk region
of Stavropol’ territory, the feeding regime of the fox changed an-
nually and seasonally according to the number of mouse-like rodents
(Table 41). A decrease in remains of secondary food species in
feces and stomachs serves as a characteristic index of food
favorability of the fox in this region. They were filled with mouse-
like rodents in the years 1937/38 and 1940/41; on the contrary,
remains of various secondary foods in seasons of decrease in num-
bers of small rodents, and the small amount of the latter in feces
and stomachs in autumn and winter 1938/39, 1939/40 and in the
summer of 1940 indicated unfavorable nutritional conditions.

In the Kushka valley (southern Turkmeniya, 1954), the fox diet
varied extraordinarily with the seasons of the year. In spring and

*Sic; the proper group name is cottontail rabbit—Sci. Ed.

354

522

Table 41. Contents of feces, stomachs and intestines of red foxes in
chernozem herb-grass steppe of the Cis-Caucasus in various years
(in % of occurrence in analysis; A.F. Chirkova and N.N. Bakeev)

Year and Winter, Winter, Autumn Spring, Autumn
season, 1937/38, 1938/39, and beginning and

number of зрипе spring winter, of summer winter
objects 1938 1939 1939/40 1940 1940/41

Food type 70 feces 310 feces, 32 feces, 491 feces, 12
4 stom- 30 stom- 2 stom- stom-
achs and achs and achs and achs and

intestines intestines intestines intestines

Mouse-like rodents 100.0 100.0 95.2 100.0 100.0
Ground squirrels — 13.0 — 8.8 —
Brown hares 12 6.5 9.5 5.9 —-
Insectivores — 6.5 — 3.0 —
Carrion, animal — 8.5 16.7 5.9 16.7
wastes
Birds 6.3 24.0 16.7 61.8 25.0
Reptiles 24 13.0 — 14.7 —
Insects 24 52.2 11.9 61.8 —
Plants 6.3 24.0 64.3 17.6 75.0
Inedible wastes — — 2.4 3.0 —
(rags and others)
Unidentified — = 2.4 — 8.4

summer, insects served as the main foods (97.7% of occurrences),
mainly termites, locust and beetles; small rodents were encountered
in 1.1% of occurrences. In autumn and winter, insects constituted
only 58.1%, but mouse-like rodents increased significantly (18.6%),
especially the Afghan vole, reptiles and carrion; hedgehogs disap-
peared, and the number of the species of passerine birds significantly
decreased (Shcherbina, 1958).

Variations in the fox diet in the Ukraine by season of the year
are considerably less (Table 42). During 7 years of investigations
in various areas, foxes consumed 118 animal and 25 plant species.
The greatest variety of food was discerned in summer and autumn
(83 and 80 species), and less in spring and winter (40 and 56
species). Mouse-like rodents remained the main food group through
all seasons of the year, constituting 97.5% of occurrences (among
the remains of all mammals), of which voles were 76.5% (Korneev,
1956).

523

355 Table 42. Main food types in diet of red fox in the Ukraine by season
of the year (in % to numbers of different data; after Korneev, 1956)

Period,
quantity Spring Summer Autumn Winter Yearly

of data 777 170 181 198 726

Food type

Mammals

Red-toothed shrews 3.3 1.7 4.4 2.0 2.9
[Зоистае]

Brown hare 7.9 14.1 3.8 Level 9.2
[Lepus europaeus]

House mouse — 4.7 6.6 12.6 6.1
[Mus musculus |

Harvest mouse — 2.3 5.3 5.0 3.3
[Micromys minutus]

Striped field mouse 14.6 2.3 9.4 5.1 8.1
[Apodemus agrarius]

Common field mouse 7.9 12.9 9.4 10.0 10.0
[A. silvaticus |

Sagebrush vole — — 4.9 5.1 2.9
[Lagurus lagurus]|

Northern red-backed 2.2 4.1 5.3 3.0 SF)
vole [Clethrionomys
rutilus |

Water vole [Arvicola 36.7 21.1 17.6 7.5 20.3
terrestris |

Field vole [Microtus 20.9 11.7 10.5 7.0 12.0
agrestis]

Common vole [M. 61.5 50.6 60.2 15.0 61.7
arvalis |

Birds

Passerines, not further 3.3 3.5 5.3 0.5 3.2
identified

Domestic fowl 0.6 7.0 I 25 2.7

Reptiles

Sand lizard И 5.8 3.3 2.0 3.2

Insects

Mole-cricket [Medvedka] 6.2 14.1 6.0 0.5 6.4

Ground beetle lee 8.8 led —- 27

Dung beetle 0.6 7.6 1.6 eS 2:7

Мау cockchafer beetle — 53 3.8 -— 2.2
[Melolontha] larvae

Plants

Sugar beet — — я 5:9 22

Pear fruits 0.6 D3 11.6 0.5 3x7

Carrion; farm animals 6.7 1.2 4.4 7.0 4.9

and dogs

524

In the Darvinsk [Darwin] preserve in the shore zone of the Rybinsk
reservoir, the fox ate more mouse-like rodents in April—June (80% of
occurrences in diet), when they were left unprotected during times of
flooding, than in August-October (72% of occurrences); in the cen-
tral part of the preserve, rodents were more often eaten in autumn
(93.7%) as compared to spring (76.9% of occurrences). Consumption
of secondary foods varied by biotopes, especially hares, ducks, am-
phibians, fishes and mollusks. The ratio of food types varied sharply
depending upon the volume of spring flooding. In winters following
heavy spring floods, when mouse-like rodents almost disappeared,
they had little significance in the fox diet, and after a spring with little
water, they constituted, on the contrary, the main food of the fox. In
the first case, the significance of secondary foods was strengthened—
hares, insectivores, all species of birds, fishes, plants, carrion, human
domestic refuse (Kaletskaya, 1957).

Variations in the diet of the fox in different biotopes of southern
Turkmeniya is very indicative (Table 43). In the Kushka valley and
in the pistachios, insects served as the main food and in the Kushka
valley, almost exclusively insects and arachnids. In desert-steppe
biotopes, small rodents stood in first place in the diet, but in years of
low numbers (1958), the fox also fed on insects and lizards, skinks
and tortoises.

Home range. The fox home range usually occupies an area of
several square kilometers. In the central forest belt of the European
part of the USSR, its diameter is equal to 6-10 km. The size and
character of the biotopes of the home range change depending upon
the season of the year, and food abundance. As a rule, during the
breeding period, especially in years of food abundance, the home
range diminishes, but in the autumn—winter period and with food
scarcity, it enlarges considerably; in years of starvation the fox moves
into biotopes not usual for it near human settlements. After dispersal
of the litter, foxes hunt and feed singly. In the case of disease or
injury, the fox always strives to reach its home range. The individual
home ranges of foxes, especially in winter time, overlap each other.
In the steppes of the Cis-Caucasus in some years, it was possible to
observe tens of foxes mousing simultaneously within one field of
vision.

In Lapland preserve, in winter, the fox develops a territory of 30
to 50 km? and more. Judging by marking [of foxes] in the forests of
Kalinin district, one fox litter one and a half years old in winter held

356

525

Table 43. Diet of foxes in various biotopes in southern Turkmeniya (Badkhyz;
spring-summer 1954 and 1958; findings in % of number of samples
investigated’; Shcherbina, 1962)

1958

1954

Pistachios Kushka Desert- Pistachios Desert-
river valley steppe steppe
Mammals 30.9 2.6 75.0 21.2 60.7
Insectivores*® 0.5 0.6 0.7 — =
Hares 0.1 — — 1.2 0.1
Mouse-like 28.2 п 130 20.0 60.6
rodents
Birds 8.1 3.4 19.0 23 9.1
Bird eggs 0:2 — — — —
Reptiles 8.0 0.8 2.4 24.7 35.8
Tortoise eggs 0.6 0.3 — 1:2 6.1
Arachnids 21.8 10.0 2.4 14.1 4.5
Insects 86.1 97.7 26.9 91.8 56.6
Myriapods 0.2 — — — —
Pistachio nuts 1.9 — —- 24.7 —
Other plants — 5.4 — — —
Carrion Я] 0.9 1.3 a —

to a total distance of 2-3 km from the burrow. In the forest-steppes,
foxes roam considerably more widely. On the left bank of the Dnepr
(Kiev district), the young foxes of one litter were caught in the course
of a summer in various burrows at distances no greater than 4—5 km
from their birth place, but in winter, not only this litter but also others
usually dispersed in a radius of 15—40 km and in individual cases
emigrated more than hundred kilometers from the burrow; young
foxes went out several tens of kilometers in the Tatar ASSR
(Nasimovich, 1948; Chirkova, 1955; A. Popov, 1956).

In the course of 24-hr period, from one bedding-down to the
next*, foxes in the Lapland preserve and in Stavropol’ territory moved
up to ten kilometers, and in severe freezes—more than 30 km
(Sabaneev, 1904). On another day, the line of their tracks sometimes
continues farther, sometimes returns backwards, repeating the previ-
ous route.

“In 1954—2780, in 1958—1275 samples (mainly feces).

382% hedgehogs.

*In the Russian original, of lezhki do lezhki; Ш. “from lying down to lying
down’—Sci. Ed.

357

526

Foxes feeding on carrion, ог on carcasses of animals killed by
wolves, sharply contract their hunting territories. They remain at the
carcass for days and sometimes weeks, and after feeding, several
individuals lie up in the nearest bog. In Lapland preserve, their home
range is no more than 5—8 km’; when resting, the foxes lie several
hundred meters from food source*, rarely up to 1%—2 km. In forest
reserves near Moscow, in the winter of 1951/52, distinguished by a
small quantity of small rodents, from 3 to 5 foxes fed on discarded
scraps of meat from a nearby slaughterhouse, and on a route of 6—7
km there in April, 20 tracks of foxes encountered roaming in the
vicinity of the food source; in January, before the trimming of the
meat, a 15-km route intersected a total of 4 tracks evenly on this
route. In the Brovarsk region of Kiev district, from 2 to 8-10 foxes
approached baits placed in six areas near dens of summer litters
(Nasimovich, 1948; A.F. Chirkova).

In coniferous forests of Manitoba in Canada, individual fox ter-
ritories have a diameter up to 19 km, but in summer are two times
smaller (Seton, 1909). In mixed and deciduous forests of the USA,
hunting territories cover tens and hundreds of kilometers (Armold and
Schofield, 1955), but in the center of Europe, marked fox cubs ranged
a distance of several hundreds of meters to ten kilometers, rarely
farther (Niethammer, 1937; see below, “Seasonal migrations and trans-
gressions’’).

Burrows and shelters. The choice of places for burrows is guided
by protection, soil and feeding factors. Red foxes prefer to construct
their burrows far from populated places, but cases are known of
burrows in immediate proximity to human habitation. Burrows are
often situated on the slopes of hills and mountains or in ravines,
bluffs and steep banks of rivers, lakes and seas, in irregular lands—
ditches, depressions, gutters, in clefts of rocky defiles, and also
neglected gullies, in places that are neglected, abandon, and usually
not flooded by high waters. Sometimes, they are restricted to parts of
forest, shrubs, or thickets of weedy grass and shrubs, to fields of tall-
stemmed cultivation—cereals, sunflowers and others, to thickets of
reeds, canes and sedges. In the forest, foxes often dig burrows among
the roots of tall-trunked trees.

In the north, the fox constructs its burrows most often on slopes,
usually in forested sections giving way to tundra, more rarely in shrub
tundra, and very rarely in the moss or lichen tundra, where it some-

*In Russian original, privad; lit. lure, bait—Sci. Ed.

527

times occupies burrows of Arctic foxes. It digs burrows in light, dry,
sandy, sandy loam, clay-loam; well-drained soils. In the central forest
zone of the European part of the USSR, the burrows of foxes reach
the subsoil layer or the lower part of the alluvial layer (Kolosov,
1935a). When excavated among the roots of trees, they can serve for
many decades, and if partly destroyed, they are rebuilt anew; in the
steppes, burrows are used for several years (A.F. Chirkova).

In deserts, fox burrows are situated on sand dunes and in saksaul
stands, in the Badkhyz preserve, often in colonies of the great gerbil
[Rhombomy opimus]. Here, the fox also uses burrows of porcupine
[Hystrix leucura], wolf and other large animals. Of 118 burrows here,
65.3% were in great gerbil colonies, 22.9% were independently dug,
7.6% in porcupine burrows and 4.2% in the clefts of rocks and ero-
sion channels (Shcherbina, 1961а, 1962). In sandy deserts, burrows
are easily destroyed.

In particular areas where there are rich food reserves with good
conditions of protection—among rocky outcrops of limestone having
natural solution chambers, or near broad stands of reeds in the open

Fig. 125. Den burrow of red fox in arable land. Voronezh district. Photograph by
A.F. Chirkova.

358

528

steppe, foxes live in colonies, constructing many tens of burrows in
а. small expanse (Cis-Caucasus, Chirkova, 1947, 1952a; Crimea, М.О.
Emel’ yanova). In the Staromar’evsk region of Stavropol’ territory, an
extent of 4 km of rocky outcrops within the hilly steppe, 40 burrows
were constructed, which were used for the cubs in various years. In the
Manych [river] steppes along heavy growths of reeds on the flood plain
of the Chograi river, up to 80 burrows were dug along an 8-km segment
of which 11 were occupied by litters during one season (Fig. 130).

In the forest-tundra, burrows are situated on southern, southwest-
ern and southeastern slopes of hills; in the steppe zone, western
slopes are preferred, and in deserts—northern slopes. The main en-
trances to the burrows are also oriented to the plane of the surface.
Temporary burrows in the deserts of southern Turkmeniya are often
dug with the opening to the south, probably in relation to the north-
erly winds that prevail here in summer.

In contrast to the burrows of Arctic foxes, badgers, marmots, and
corsac fox, which are sometimes occupied by foxes, the burrows that

Fig. 126. Entrance to a burrow of a single male; its feces in front of the burrow.
Voronezh preserve. Photograph by A.F. Chirkova.

359

529

they excavate are usually not characterized by great complexity (Fig.
131). They are divided into den and temporary burrows; the latter
consist only of one short passage or small cave dug for concealment.
Den burrow construction is somewhat more complicated: the main
entrance leads downwards (at 40-45°), broadening to form a den,
from which passageway side-tunnels branch in various directions the
locations of which vary greatly. In the forest-steppe of Kiev district,
along the passage of a burrow, there are from 1 to 3 widenings
layered with grass or leaves. In Kolyma, the nesting chamber was 35
cm high and 50 cm in diameter. In the Ukrainian steppes and in the
desert of northern Pri-Aral and southern Turkmeniya, burrows have
up to 3—5 entrances, but those located in colonies of gerbils—up to
15; in Ust’yurt, there are often 2 entrances, and in the forest around
Vychegda, up to 3 entrances. In the steppes of the Manych [river],
individual den burrows of foxes possessed up to 32 entrances; appar-
ently, the red foxes occupied the burrow of a corsac fox (Kostin,
1956; Shilova, 1960; Shcherbina, 1961; A.I. Zubov, A.F. Chirkova).
In the Far North, from 2 to 10 entrances are known (lokhel’son,
1898; Sokol’nikov, 1927; Portenko, 1941); apparently, the burrows
with a large number of entrances belonged to Arctic foxes.

Depth of the burrows ranges from 0.5 to 2.5 m; usually they do
not extend to ground water. Length of the main passage on a straight
line reaches 17 m, standing at 5 to 7 m on the average (Kiev district,
Stavropol’, Badkhyz preserve). Diameter of the inner passages of the

- burrow in steppe soil is 15 to 20 cm in height and 25 to 30 cm in

width; in desert soils the dimensions of the burrow are larger; width
of the main entrance is 30 x 40 cm and more, usually rounded or oval
in section, sometimes of nearly slit-like form, its height—up to 70
cm.

In spring, during the cleaning of the burrow, the fox, with strong,
rapid movements, at first with front paws, and then with a kicking
movement of the hind feet, throws the discarded soil a distance of 2—
2.5 m. After the appearance of the young in the burrow, this soil is
trampled, forming a spot on which the fox cubs play and to which the
parents bring food. The general area of the den burrow is from 12 to
72 m’, on average—40 пл? (Kiev district); in the deserts of southern
Turkmeniya burrows with areas from 7.4 to 52.8 m? are known
(Shcherbina, 1961a; A.F. Chirkova). Rarely, red foxes litter outside
a burrow, in a hollow or near the fallen trunk of a tree, in a pile of
firewood logs in the forest, etc.

530

Fig. 127. Entrance to the den burrow of a Turkmeniyan red fox, built in a great
gerbil [Rhombomys opimus] colony. Badkhyz preserve, southern Turkmeniya.
Photograph by Е.1. Shcherbina.

360 When the young animals begin to grow up, foxes usually change
burrows due to parasitic infestation (Kiev district, Badkhyz preserve).
Daily activity and behavior. Red fox activity depends mainly

upon feeding conditions. In places where the fox is fairly free from
pursuit, it may be seen mousing at any time of day or night. Preferred
hours of hunting are early morning before sunrise and late evening—
beginning of night. The fox spends a longer or shorter time in hunting

—

531

depending upon the abundance of food. Often, it rests in the hot
midday hours and, apparently, for a short period in the dark time of
night. When the foxes go in search of food, the fox cubs come out
of the burrow and play, but hide themselves at the slightest danger;
when the mother returns, the cubs suckle and feed on the food pro-
vided, after which they play for a while and go into the burrow for
a rest. In the evening, this picture is repeated.

At the end of summer, after the litter leaves the burrow, the
animals experience a wandering life in search of food, at first to-
gether with the mother, lying down to rest in one or another burrow.
In autumn, after the final dispersal of the litter, and later-—in winter—
the young foxes scatter and feed singly. By the time of first snowfall,
the young foxes are inactive for two-three days and lie up anywhere
in a protected place. In autumn and winter, the adult foxes and the
growing young spend almost all the time in the open air, resting in
forms between feeding hours. The young foxes often retreat into
burrows, usually in wet weather or hiding from danger. Both adult
and young foxes in the middle regions [of the country] in the very
severe frost (-30°C and below) usually spend the day in burrows
(V.G. Heptner).

The red fox feeds on carrion only in the late hours of the evening
and in night, and leaves at sunrise. If the carrion is visited by wolves,
the fox follows after them early in the morning.

In the second half of winter, with the onset of estrus, one can see
“wedded” foxes in midday. In the park on Pogonno-Losinoe Island in
Moscow, and in Byelorussia, four males happened to be observed
running after one female (Serzhanin, 1955; A.F. Chirkova), and in the
steppes of Voronezh district (Ognev and Borob’ev, 1924)—up to 10
males, who yelp and fight among themselves. At the time of rut,
foxes are very excited and almost never rest.

After mating, the female cleans the burrow and leads a more
secretive way of life; at the end of pregnancy she spends most of the
time in the burrow. In the first days after birth of the cubs the mother
does not leave the burrow, and the male brings her food. After that,
the male mostly avoids the litter and lives alone, occupying one or
another empty burrow; some males do not accept sharing in the
feeding of the fox cubs.

The daily cycle of foxes varied geographically, depending chiefly
on differences in main foods (nocturnal, diurnal, crepusular animals)
and the climatic factors (temperature, wind force, precipitation), as

362

360

532

well as upon human activity. Thus, in some places in the deserts,
where it is not especially persecuted, it fears man very little. In the
beginning of the 40’s in the Badkhyz preserve “the behavior of foxes
takes sometimes the form of such ‘trustfulness’ as is never observed,
for example, in the European part of the Soviet Union. In June, one
fox visited the site of the expedition for several successive days,
either in the evening or at dawn, and coming within a distance of
100—150 meters, barking sometimes for half an hour without a pause;
it was not even disturbed by activity which prevailed in the camp
during formation of the caravan composed of eleven pack animals. At
night, foxes traveled along their paths passing within 10-15 m of
sleeping people. In the 30’s, in the Kushka valley they stole the
provisions from tractor-drivers in camp. Sometimes in the daytime
foxes approached to within 100—150 т of wells and left only when
frightened” (Heptner, 1956).

Fig. 128. A litter of Turkmeniyan desert foxes, Vulpes vulpes flavescens Gray, by
a burrow in the open desert, A badran trunk, Ferrula badrakema, is nearby.
Badkhyz preserve, southern Turkmeniya. Photograph by Yu.K. Gorelov.

535

At the northern border of the forest—in the Lapland preserve
(Nasimovich, 1948), the red fox in summer is more active at night
while in the winter time—in the morning. Here, foxes go hunting
1-2 times per day/night, and hunt with short pauses. In the taiga of
the Fast East, foxes are very cautious and feed in the evening or early
morning (Slyunin, 1900).

In the steppes of the Cis-Caucasus, foxes always hunt in day-
time, but, only in the period of reproduction; the adults do this more

361 Fig. 129. Old den burrow of the Turkmeniyan fox in pistachio grove in the
mountains of Gyaz’-Gyadyk. Badkhyz preserve, southern Turkmeniya. Photograph
by E.I. Shcherbina.

534

2 3km
(Ee ee НЫ]

362 Fig. 130. Section with a high concentration of fox burrows in floodplain of Chograi
river, overgrown with reed. Virgin steppe along the East Manych [river],
Stavropol’ Territory (from Chirkova, 1947): 1—Hills; 2—Farms, sheep-folds;
3—Fox burrows without litters; 4—Burrows with litters, 1938; 5—Burrows with
litters, 1939; 6—Burrows in which there were litters both years; 7—Burrow of

corsac fox with litter, 1939; 8—Flood plain border.

stealthily, and at the end of summer and in autumn, inexperienced
young foxes are often encountered approaching humans at close range.
In the desert in summer, the fox often never goes abroad at all in
daytime, protecting itself in the burrow from sunstroke. At the same
time, even in the very hottest time of year, for example in the Badkhyz
desert, foxes sometimes continue to hunt until 8-9 o’clock in the
morning and later, when it is already hot and the sun shines strongly,
and in the evening, it hunts long before the sun sets, when the heat

363

535

‘has only started to decrease. This is, apparently, connected with the

fact that one of the chief food items of the fox—the great gerbil—
is a diurnal animal. Besides the mentioned period, and especially in
winter, foxes in Turkmeniya are, to a considerable degree, active
during daytime. This is partially explained in that several rodents
serving them as prey, for instance red-tailed gerbil [Meriones
erythroura], are active in summer mainly at night, but in the period
when it is not hot, turn to a diurnal mode of life (V.G. Heptner). In
general, foxes in nature are distinguished by broad individual vari-
ability in behavior and daily rhythm.

Seasonal migrations and transgressions. More or less regular
seasonal movements of foxes are known for the northern margins of
the [Soviet] Union, the deserts, and the high montane regions. In the
extreme North, after a mass appearance of small rodents, there is an

Fig. 131. Scheme of construction of fox burrows (A.F. Chirkova). Steppes of the

Cis-Caucasus: 1—in sandy meadow; 2—in water meadow (with litter). Forest-

steppe of Kiev district: 3, 5, 6, 8, 11, 12, 13—among arable land or winter rye (with

litters), 4, 7, 9, 10—in forest or shrubs (with litters, except 10). 4, 6, 8—former

badger burrows. Figures near nest or at separate parts designate depth under
surface: 1—exit openings, 2—main exit, 3—nest (see legend).

364

536

increase in number of foxes, which immigrate to the tundra searching
for food. The farthest known transgression of foxes outside the breed-
ing zone is to Novaya Zemlya, the coast of Kara Sea northeast of the
Bol’shezemel’sk tundra, the northern extreme of Yamal, northern
Gydansk Peninsula, Dixson Island, the central part of the Taimyr
Peninsula to 72.5°N. Lat., Lyakhovsk Island below 73—74°N. Lat.
and Wrangel’ Island, 300—700 km from places of reproduction of the
species (Middendorf, 1869; Zhitkov, 1913; Mineev, 1935; Heptner,
1936; M. Popov, 1953; N.V. Provorov). In addition to meridional [N—
$] migrations, latitudinal [E-W] migrations also exist—for example,
as in the Arctic fox, from the lower Pechora to Malozemel’sk tundra
and from Timansk tundra to Kanin Peninsula (Semenov, 1936; A.F.
Chirkova). In Kholmogorsk region of Arkhangel’sk district, a red fox
was killed which had been marked 600 km to the northeast in
Malozemel’sk tundra (Parovshchikov, 1956).

In the course of the last decades, colonization by foxes has been
taking place in the forest-tundra and southern tundra. Its frequent
transgressions and, sometimes, breeding is observed in the entire
European tundra belt, with the exception of the extreme northeast.
Advance to the north was also observed in Yakutiya and in Magadan
district (Skrobov, 1958, 1960, 1960a: Shustov and Belozorov, 1959).

With deep snowfall in the forest zone, migration of foxes into the
floodlands of large rivers occurs, and in Kamchatka and Sakhalin—
to the seashores.

In the forest-steppe of Kiev district and the Tatar ASSR, by
marking on paths, it was shown that over 1—2 years, the majority of
foxes dispersed a distance of 15—30 km, some of them, 2—5 km and
only one in Kiev district was caught 120 km from the place of
release. Emigration of foxes in the course of a year for several tens
of kilometers is also confirmed by recording of transgressions of
individuals ill with mange (Chirkova, 1955, 1957; A. Popov, 1956).

In southern Turkmeniya—Badkhyz preserve—foxes in winter
spread more or less uniformly over the territory; at this time, waterholes
do not play any role in the animal’s lives. In summer time, inhabi-
tation deep in the desert becomes difficult for it due to the distances
to be traveled to water. In this connection, in the hot time of year,
foxes, if they do not abandon these regions of the desert completely,
then in all events their habitation here is notably reduced. Movements
take place in two directions. The population of Badkhyz foxes, on the
one hand, gathers on the periphery—in the valleys of Kushka,

365

537

Egri-Gek and Tedzhen rivers, where some small pools of water re-
main, and on the other, they more or less concentrate near the wells.
Water remaining in wells from the watering of sheep is used at night
by foxes (Heptner, 1956).

In the mountains of the Caucasus and Trans-Caucasus, Middle
Asia and the Altai, in September—October, with the falling of deep
snow, foxes descend from the alpine and subalpine zones to the forest
belt and lower. In the Sayans they wander down to less forested
places where the snow is not so deep. The movement of foxes from
the high mountains to the lower forest belt takes place also in
Kamchatka. The reverse movement of foxes to the mountains in
spring is less obvious.

Reproduction. The red fox reproduces once per year, in spring.
Some females become capable of reproduction at the age of 9-10
months and give birth to young at the age of one year.

All females become sexually mature at the end of the second year
of life.

In the middle latitudes, the preparation for estrus in females begins
about two months before it starts, in the majority of cases in Decem-
ber. The organs of reproduction proliferate in a definite sequence,
changing internally and increasing in dimensions. At first, the neck
of the uterus swells. The vagina thickens by almost three times, its
inner epithelial layer becomes cornified, which is characteristic for
the beginning of estrus; in the mucous secretions of the female, clumps
of exfoliated epithelium are observed. Obvious growth of the uterine
horns begins about two weeks before estrus; at the time of estrus, the
uterine horn increases almost double, becoming resilient and elastic.
At the time of estrus, the ovaries increase 1.5—2 times; follicles in it
mature, forming Graffian vesicles. Ovulation in the fox is spontane-
ous; 1.е. not dependent on copulation. The mature follicles rupture not
simultaneously and in their places corpora lutea appear, in connec-
tion with which fertilization is possible at various times by [different]
copulatory acts. At the moment of estrus, the sexual organs of
females attain the greatest degree of swelling and elasticity.

In nature, the sexual cycle in males begins in August-September,
when the process of sperm formation begins, causing swelling of the
testes. The testes attain their greatest weight in December—February
when the spermatozoids mature; at that time, sperm appears in great
quantity in the seminiferous tubules and in the epidydymus (Chirkova,
1941, 1947, 1954, 1960) (Fig. 132). In the silvery-black fox in

1937 1938 1939

Fig. 132. Sexual cycle of the male red fox (by testes weight) in successive

years with various feeding conditions (from Chirkova, 1947): 1—average

weight; 2—maximum weight; 3—minimal weight; 4—presumed path of curve
(no materials).

captivity, the appearance of prostate gland secretion, essential for
insemination, is delayed by 1.5—2 weeks as compared to the testes.
The female permits the male to copulate for a period of several
days; in this time, the mammary glands of the female swell to 0.5 cm
in diameter (Zaleker, 1937, 1939; Starkov, 1937; Il’ina, 1952; Kler,
1953). In fur farming, cases of repeated copulation are known, which
occur when fertilization does not take place the first time, after abor-
tion or death of the new-born cubs. Copulation is accompanied by a
copulatory tie between male and female which sometimes continues
366 for more than an hour. After mating, the fertilized eggs are implanted
in the horns of the uterus on the 8th—10th day. Pregnancy in foxes
lasts from 49 to 58 days, often 52 days.
Foxes often copulate in burrows (Cherkasov, 1867 and others).
Up to 13“ fox cubs may occur, on the average, 4—6. The lactation
period lasts 1% months. The young foxes remain in the burrow about
1.5—2 months. Playing and searching for food, they gradually begin
to go farther from the burrow, and finally do not return to it any
more, leaving together with the mother. Already in August, in middle
latitudes, young foxes begin to lead a fully independent life; in

“Up to 14, according to unverified data.

365

539

% of females

J ь
3 а
1-10 11-20 2-91 1-0 1-7 21-91 1-40 1-20 2-28 50 10-20 21-31 140
December January February March April:

Fig. 133. The course of estrus in the red fox of European part of the USSR for two

successive years (1938—1939) with different feeding conditions. Percent of females

in estrus to the total number of examined females. Materials from 10
steppe, forest-steppe and taiga districts (from Chirkova, 1947).

southern steppes, this takes place in July and in the years of poor food
conditions—in June.

Reproduction of the red fox varies both geographically and by
year (Figs. 133, 134); the time of reproduction and fertility both vary
(Chirkova, 1960, 1961). Reproduction in the desert and steppe zones
occurs earliest of all, while in the taiga and forest-tundra zones, it is
significantly later. In the southern steppes, estrus in the female red
fox occurs most often in December—February, in the forest zone of
the European part of the USSR, in February—March, and in the forest-
tundra zone, they come in heat in March—April. The testes of males
attain their maximal weight in the southern steppes in December, in
the northern forest-steppe—in February, and in the forest zone—
February—March. Within the whole forest zone of the USSR, rut of
foxes continues for 5 months. Separate cases of rut in all zones end
usually in April.

Fox cubs appear at the burrows mainly 2'/, to 3 months after the
beginning of estrus in females. In the forest zone of the European part
of the USSR, whelping occurs chiefly in April, and fox cubs appear
near the burrows in the course of May—June. In the steppe zone, the
fox gives birth to young primarily in March—April, and the young
animals appear near the burrows in April-May. Litters are found at

367

366

540

the burrows in the entire territory of the steppe and forest-steppe
zones, and probably, in the forest zone, during about 5 months.

In the deserts and semideserts, red fox reproduction is greatly
prolonged. In Badkhyz preserve (southern Turkmeniya) the first signs
of the onset of nuptial times are observed in November, when the
foxes begin to move in pairs. Especially intensive mating is observed
in December and comes to an end January. Whelping takes place in
February—March. The first early litters were observed in the middle
of February (Shcherbina, 1958). Here, fox cubs are met with at the
burrows during the course of 7 months. In montane regions, judging
by separate observations reproduction takes place in the various belts
at different times; in the lower, it starts nearly simultaneously with
the surrounding plains, while in the higher it is delayed.

The appearance near the burrows of the first litters of fox cubs
in the deserts is sometimes noted already in January, in the steppes
and some montane regions of the south, from January—February, in
the forest zone from March—April, and in the northern forest zone
and in the forest-tundra, from May and later. All times of reproduc-
tion are, on the average, delayed in a northeasterly direction both for
the territory of the Union as a whole, and in each natural zone. In the
territory of the entire Soviet Union, the time of appearance of litters
of fox cubs on the surface extends for half a year (Fig. 135).

хп Ш лм
А В

Fig. 134. Variation in the sexual cycle of the red fox т various natural zones of the

European part of the [Soviet] Union (from materials of a series of years, from

Chirkova, 1960). A—Estrus in females according to the laboratory determinations

in % of the total number of examined specimens; B—Weight of male testes : 1—

southern steppes; 2—northern forest-steppe, mixed forests and southern taiga;
3—northern taiga. Months are plotted on the abscissa.

Fig. 135. The time of appearance of the earliest litters of fox cubs at the surface
near burrows. Roman numerals indicate months (from Chirkova, 1960).

Fertility of the red fox is also geographically variable. The least
number of young—an average of 3.5 up to 6—is observed in the
Karelian ASSR; an average of 4.9 up to 8—in the deserts of Turkmen
and Uzbek SSR, and up to 8—9— т high montane zones of the south.
In the zone of the southern taiga and broad-leaved forests of the
European territory of the Union, in the southern regions of Krasnoyarsk
territory and in Buryat ASSR, a fox family has an average of 4.0-4.9
young. There is higher fertility in the Stavropol’ steppes and western
Siberia—5.0—5.6 on the average and up to 13 cubs (А.Е. Chirkova).
Fertility is high in the northern forest-steppe of Volzhsko-Kamsk
territory: 5.8 on the average, maximum 13 (V.A. Popov).

There is great variation in time of reproduction of the red fox in
one place in various years related to variation in food conditions (Fig.
133). Strong frosts and snowstorms delay estrus. The individual times
of estrus and rut vary strongly depending upon age, growth condi-
tions of the individuals in various fox generations, upon the course of
female reproduction in preceding seasons, etc. Thus, in Kanino-
Timansk tundra the time of rut in successive years differed by 1-5
months. Deviations in the time of beginning of estrus in the forest
zone of the European territory of the USSR for a 3-year period
constituted 3% months. Judging by study of the reproductive organs
of foxes obtained in winter, females in estrus condition might have

368

369

542

been found in this zone starting from January and even the end of
December up to the middle of April. During successive years, the
beginning of estrus was offset by 1-1.5 months. There were strong
changes in intensiveness of rut. In favorable years, concerted estrus
took place in the course of 1 month; in the unfavorable years, it
expanded to 2% months. In the Stavropol’ steppes, the time of estrus
is displaced in successive years by 2—3 months. The whole period of
reproduction of red fox in each place from the beginning of rut to the
appearance of fox cubs at the burrow is about 6 months, but even
longer in the desert zone.

Fertility also changes annually; primarily, the quantity of repro-
ducing males and females varies (Fig. 136). In years of good nutrition,
70-80% and up to 100% of females participate in reproduction; at
that time, all den burrows are occupied and individual foxes deliver
the young even outside them.

In an unfavorable year, up to 50-70% of females remain barren
(did not come into estrus, not covered by males, with resorbed em-
bryos and also aborted ones). The majority of den burrows at that
time were empty. As a result, reproductive increase sharply decreased
compared to the preceding year.

Reproduction of the various subspecies of red fox, especially in
Siberia and montane places, has not been studied.

In Bulgaria, foxes reproduce at various times also. The rut of
foxes in southern Bulgaria begins in January, and in northern Bul-
garia—in February. The female fox gives birth to 4 to 9 and up to 12
young. In the USA the red fox gives birth to up to 10 cubs; on the
average 5—6; in the eastern states, they are born at the end of March—
the beginning of April; in the steppes of Oklahoma there are 3—9 cubs
(Selko, Krefting and Fletcher, 1942; Hamilton, 1943; Trippensee,
1953; Atanasov, 1958).

Growth, development and molt’. Cubs are born blind, with closed
ear pinnae and without teeth. Their body is covered with dark-brown
fluffy fur. The tail is narrow at the tip, and in the majority of cases
with fluffy white fur. The weight of the newborn wild fox cub is 56—
110 g, averaging 86.9 g, on fur farms, 50—150 g, normally 80—100 g,
body length about 14.5 cm, tail 7.5 cm. The cub’s eyes open on the

чп wild foxes, this aspect is insufficiently studied; therefore, separate indices
are adopted from the practice of fur farming (Starkov, 1932; Bogolyubskii, 1939;
Leshchinskaya, 1950; Il’ina, 1952; Eremeeva, 1954).

368

543

— == 7

@-:
°, Scale of
100 Moscow district Steppes of Cis-Caucasus nee
8
7
6
5
4
3
2

Igavrriugvriruy

ol Wel Tee Vel eae

1937 1936 1939 1937 1938 1939
. Arkhangel'sk district Kirov district Scale of
100 abundance
90 9
60 8
70 7
60 6
20 я
40 4
30 3
20 2
10
ПТ ЛИШ ДДД I
IGS? 1938 1939 19ST, 1938 1939

Fig. 136. Red fox reproduction in relation to the number of small rodents in
various natural zones of the European part of the USSR (from Chirkova,
1947). 1—Number of small rodents in scale of abundance; 2—Percentage of
females ready for reproduction from total number of investigated animals
(laboratory analysis). On the abscissa, years divided into quarters.

544

13th—15th day on farms, at that time also the ear canal opens and
teeth appear in the upper jaw and after about 3—4 days more, in the
lower. At the age of 3—4 weeks, fox cubs in nature begin to come out
from the burrow and are gradually schooled in different kinds of
food, which is brought by the parents. The lactation period lasts about
6-7 weeks. The fox cubs are born short-legged, large-headed and
with broad chest. Later, the extremities grow intensively, and in 3—
4 months, the whelps are long-legged, narrow-chested and sinewy.
Body proportions of the adult animals are attained at the age of 6—
7 months.

Replacement of milk teeth by permanent ones in southern Ukraine
takes place before the age of 4 months*; in the middle of July, the
dental formula of the young animals is the same as that of adults.
Later, until age of maturity, only the canines obviously grow, and to
the end of life, wear more quickly than other teeth. In contrast to the
Arctic fox, tooth wear in young red foxes begins with the incisors of
the lower jaw and with aging, it occurs in a definite succession. On
this basis, it is possible to determine the age of foxes according to the
condition of the teeth (Brauner, 1914; Grigor’ev and Popov, 1940,
1952; Smimov, 1960).

In Kiev district, weight of wild fox cubs at about 1.5 months old
is from 900 to 1350 g with body length of 36 cm, and the tail
13.5 cm; at 2.5 months—from 2 to 2.5 kg*’ with body length of
61—62 cm and tail 26-30 cm. In adult foxes and yearlings caught
from November to February in the forest-steppe and steppe zone
(especially the former), average body length reaches 72.1 cm and up
to 89 ст, the weight of a carcass without skin averages 4.4 kg,
reaching 8 Ко” (А.Е. Chirkova).

The life duration of the red fox in captivity is up to 25 years. In
nature, foxes live for only a few years, usually not more than six.

The development of the pelage was studied in the silvery-black
foxes on farms near Moscow (Leshchinskaya, 1950; Eremeeva, 1954).

46On farms, the replacement of incisors in the fox takes place on the 90th—105th
day; after this, during the course of up to 150 days, the canines and premolars are
replaced.

47On farms the average weight of a month-old fox cub is 700—750 g and after
that the monthly weight increment is about | kg.

‘8Utilizing measurements of more than 2,000 foxes from all zones of the Soviet
Union.

“The fresh skin of a red fox weights 500-600 g.

370

545

The formation of hairs already proceeds in embryos and in them the
first hairs appear on the nape on the 415 day of pregnancy. The
shedding of hair of the first coat takes place in foxes beginning with
the 35th and up to the 75th—80th day of life; the guard hairs are lost
first, the underfur by the 110th—120th day, in August-beginning of
September. Intensive shedding of guard hairs of the juvenile coat is
observed in the second half of August, irrespective of the time of
birth. In August, vigorous development of the hairs of the first winter
(adult) coat begins; its main mass matures at the beginning of Decem-
ber.

In the adult silver-black foxes, molt takes place twice a year. The
spring begins in February, and complete shedding of winter pelage
occurs from March until July; growth of the new proceeds simulta-
neously. Pelage replacement begins in the pregnant females, after that
in barren ones, then males, and last of all in old individuals. Early
growth of the winter hair becomes obvious from August on, and
proceeds until the beginning of November, when it ends in full de-
velopment of the winter fur. Maturation of the fur in a warm autumn
is delayed for 1 to 2 weeks; with an early frost, it is accelerated.

In nature, the timing of molt and of juvenile hair in the red fox
change depending upon latitude and longitude of the location, nutri-
tional and meteorological conditions, sex, age and condition of health.
The more northern and eastern the place, the later molt begins in
spring, and the earlier the maturation of winter fur occurs. In the
temperate zone (in the plains), molt of the fox begins in February—
March, but in the southern deserts, even earlier, proceeding vigorously
in April—May and lasting until the middle of summer; in the north of
western Siberia (in Yamalo-Nenetsk Nats. Region), from end of
March—beginning of April. In montane regions, it is delayed; in Ar-
menia, molting foxes were repeatedly observed in July and even at
the beginning of August (Dal’, 1944, 1948b, 1949; Rakhmanin, 1959).
Growth of winter fur in the deserts begins in July, and by December
foxes are clothed in dense winter fur (Turkmeniya; Shcherbina, 1961а).
Maturation of the winter fur of the red fox in other zones takes place
in the course of November and December. In different years, incom-
plete pelage growth is observed in January and even later; some foxes
in these years simply never acquire complete winter fur.

Enemies, diseases, parasites, mortality and competitors. The
enemies of the red fox are not numerous. Of mammals, the wolf
represents the greatest danger, lynx and wolverine considerably less,

546

and among birds, the large raptors—eagles and sea-eagles (about 1%
occurrence in castings; Gusev and Chueva, 1951; A. Popov, 1954).
In the Cis-Caucasus, the imperial eagle and fox mutually harm each
other—the eagle sometimes preys on the fox and the fox destroys its
eggs.

The majority of diseases of the fox have not been studied in
nature. Nonetheless, from time to time disease breaks out among
foxes, causing massive mortality. They are connected with deteriora-
tion of feeding conditions, mainly disappearance of small rodents.
Food scarcity is accompanied by the emaciation of mature adults and
promotes the appearance of outbreaks of various diseases. Some red
fox diseases are, probably, common to the small rodents on which
this animal feeds, as well as to other animals. Disease transmission
is favored by high density of the fox population and the behavior of
hungry animals (frequent biting of one other, cannibalism).

High mortality among foxes was recorded during the mid-30’s,
40’s, and mid-50’s of our century. The plague is expressed most
sharply in places with high population density—in the steppe and
semidesert zones of the European part of the USSR. Individual hunt-
ers found there tens of fox carcasses, and many diseased foxes. An
especially severe epizootic and mass mortality of foxes appeared in
1944—1947 in the European territory of the USSR, in the Caucasus
and in Kamchatka, and to a lesser extent—in Yakutiya, western Si-
beria and the Altai, when mass reproduction of small rodents, followed
by their disappearance led to starvation and the outbreak of disease
among red foxes. In silver-black foxes on fur farms, a great number
of diseases were revealed. It is necessary to realize that wild foxes
are included in the chain of many infectious diseases of animals and
human.

Among wild foxes in the USSR, rabies is widespread, occurring
in all natural zones and especially often appearing in the mid 1940’s
in the territory of the European part of the USSR. The disease also
was recorded in many foreign countries. In the USA, of hundreds of
foxes caught in an infected region, the infection rate of rabies was 3.1
to 5.8%. In the extreme north of Arkhangel’sk district, carnivore
distemper—a neuroviral disease—manifested itself among foxes in
extreme northeastern Siberia, and particularly sharply in Kamchatka;
it already was known early in the first quarter of the past century. In
the tundras of Arkhangel’sk district, the Arctic fox is the main virus
vector. In years of outbreaks, up to 20% of the red foxes are virus
vectors.

372

373

547

Carnivore plague appeared chiefly in the steppe zone and some-
times caused general epidemics. In individual cases, epizootic jaundice
(leptospirosis) and tularemia were revealed, but foxes are not very
susceptible to becoming ill with the latter. Foxes may also become
sick with listerellosis and spirochetosis, and act as vectors in the
spread of erysipelas infection, brucellosis and tick-born encephalitis
(Sokol’nikov, 1927; Chirkova, 1941, 1952; Isakov, 1949; Pavlov,
1953; Vyshelesskii, 1954; Dunaeva, 1954; Sludskui, 1954; Kantorovich,
1956, 1956a, 1963; Rementsova et al., 1956; Galuzo, 1957;
Karochevskaya, 1958; Tirkel’, 1958; Bindrich et al., 1959; Wood
[and] Davis, 1959; Ismagilov, 1961; Morozov, 1961).

A disease of unknown etiology and mortality among foxes and
other carnivores took place in the region of Lake Sartlan in Novosibirsk
district, due, as was supposed to poisoning by toxic substances leach-
ing from the ground, but this was also disputed (Berman and
Strusevich, 1957). The possibility was considered of it being due to
an acute form of encephalomyelitis which was observed in silver-
black foxes (especially young) in nurseries (Isakov, 1945; Vyshelesskii,
1954). In individual cases, the infection of foxes with plague (Bacil-
lus pestis; Rall’, 1958) was observed. In some years the foxes are
found to be infected with large quantities of fleas and ticks, especially
in the desert and steppe zones; many species of these ectoparasites
serve as Carriers or transmitters of dangerous infections of humans
and domestic animals.

The itch mite Sarcoptes scabiei induces mange in foxes. In the
extreme form in which this disease appears, the body of the fox
becomes almost naked and covered with hemorrhagic scabs; such
animals gradually die. Mange on foxes in the USSR has been known
since 1919 (Chirkova, 1957). It first appeared in the form of several
foci in the southwestern Ukraine, and then spread over a vast territory
(Fig. 138). At the present time mange is established in foxes of the
steppe, forest-steppe and southern forest belts of the European part of
the USSR eastward to the Volga and even beyond. The spread of
mange is favored by temperate, warm and humid climate; fox bur-
rows in the above-mentioned zones are a very favorable microclimate
where itch mites and their eggs are preserved for a long time. Evi-
dence of disease and death of red foxes from mange in the forest-steppe
and steppe belts of the European part of the USSR comprise 55% of
all known cases of disease and death among foxes of these zones,
averaged over many years.

548

Oe Denne rest a eee re
1944 194.5 1946 1947 1948 1949 1950 1951 1952 1953 1954

371

549

Endoparasites of wild red foxes have not been sufficiently
studied (Petrov, 1941, 1958; Chirkova, 1948; Evdokimova, 1954;
Troitskaya, 1955; I. Romanov, 1956). On fur farms of the USSR, up
to 60, and in nature more than 20 helminth species are found in red
foxes and several species of coccidians of the genera /zospora and
Eimeria. The richest helminth fauna of foxes is in the forest-steppe
and the northern belt of the steppe zone, while the poorest is in the
desert zone. Infection in the fox population is almost 100% every-
where; only in some places and in some years does it decrease.

The nematodes most frequently found in wild foxes are Toxocara
canis, Toxascaris leonina and Uncinaria stenocephala, found in the
gastrointestinal tract, and Thominx aerophilus and Crenosoma vulpis—
in the lungs. Among trematodes, Alaria alata is most often found,
among cestodes, Taenia pisiformis and Mesocestoides lineatus; and
of acanthocephalans, Macrocanthorhynchus catulinus—all in the in-
testinal tract. The degree of infection with various species of helminths
changes not only by territory but also annually. Species specificity
has been recorded for some zones. The alveolar echinococcus,
Alveococcus multilocularis, and the trichinellid Trichinella spiralis
are especially dangerous, as is Dracunculus medinensis, the liver
fluke (Opistorchis felineus) and certain tapeworm species (Diphylidium
caninum and Diphyllobothrium latum); they are transmitted to hu-
mans, and some species are also characteristic of domestic animals.

Echinococcosis and trichinellosis are widely distributed infec-
tions among wild foxes. Echinococcus infection is known in red
foxes of ali natural zones of the Soviet Union. High infection rates
were observed in Alma-Atinsk district (25% of carcasses examined),
in Kamchatka (30.5%), in Omsk district (up to 35%) and in
Krasnoyarsk territory (average of 55.9%, and in some steppe regions,
up to 70% in individual months) (Kadenatsii, 1953, 1959; I. Romanov,
1958; Kozlov, 1961, 1962; Arslanova, 1962). In different cantons of
Switzerland and northward, 36-40% of foxes are infected with echi-
nococcus (Vogel, 1955, 1960).*

*In Russian original, misspelled Fogel’—Sci. Ed.

Fig. 137. Findings of sick and dead red foxes in various parts of the steppe zone as

an index of the general intensity and course of epizootics by years. Relative

number of sick animals and carcasses are given according to information of

correspondents (А.Е. Chirkova). |—Ukraine and Cis-Caucasus; 2—Volga region;
3—North Kazakhstan.

| |

Ht |
Z

(1
0)
$

SS \

Oe: ИД: WW feacs
Ев (55) 6 /: #8 ет

200 0 200 400 600km
eS a ee eee

372 Fig. 138. Distribution of itch mite-caused mange in red fox for 1919-1954 (from
Chirkova, 1957, simplified). |—Primary foci of mange in 1919—1922; 2— infected
area in 1923—1933: 3—-same in 1934—1940; 4—same in 1945—1949; 5— зате in
1950—1954: 6—Points of disease isolated from general area of the epizootic; the
year is indicated in circles; 7—southern boundaries of taiga forest; 8—boundary

between steppes and semideserts.

374

551

Trichinellosis in foxes is known from individual cases of infec-
tion in Omsk district, 3.8% in Stavropol’, up to 11.1-21.7% in
Khabarovsk territory and Kamchatka, 12.3% in Tatariya, from 0 to
12.5% (in various years) in the northern forest areas of the European
part of the Soviet Union, and up to 20.8[%] in the central forest
regions, 36.4% in Saratov district, 42-54% in Belovezhsk: Forest,
56.5% in the Crimea, and up to 74.7% in Bashkiriya (Kadenatsii,
1953, 1957; Mashirov, 1955; ТгегуаКоуа, 1956; Belyaeva, 1957;
Kozlov and Kontrimavichus, 1961 (1962); A.F. Chirkova).

Many helminth species common to other carnivores are found in
foxes, in particular, with domestic animals, a fact which is exempli-
fied in the Crimean fox (Table 44).

Among other important mortality factors of red foxes are forest
fires, burning in steppes and in reed thickets, heavy downpours in the
south, especially in the mountains, deep snowfall in deserts and
mountains, flooding on flood plains of large rivers, and the accumu-
lation of oil in the regions of oil extraction. Mortality due to elemental
calamities was observed in 2 to 16% of all cases, occurring particu-
larly often in deserts. Foxes also die from poisoned bait scattered for
control of ‘small rodents, as was recorded in 414% of cases, more
often in the forest zone and in the mountains.

Competitors of foxes for food are all species of small carnivores,
especially those belonging to the family Mustelidae and species of the
family Canidae. The red fox sometimes occupies burrows of the
Arctic fox. However, here the significance of foxes as competitors
(Skrobov, 1958, 1960, 1960a) is greatly exaggerated, since the fox in
the denning season selects biotopes which are not characteristic for
Arctic fox, mainly forested river banks with drier and lighter soils.
The red fox, in contrast to the Arctic fox, prefers to feed on rodents
of the genus Microtus. In the zone of contact of both species in the
tundra, the red fox is not numerous.

In forest and steppe zones, the fox competes with the badger,
whose burrows it often occupies; in steppes, the fox occasionally
lives in the burrows of corsac fox and marmot, and in the desert—
in wolf and porcupine [burrows] and in colonies of gerbils; in the
Caucasus mountains, the burrows of foxes are occupied by the forest
wild cat Felis silvestris; in Tadzhikistan, the steppe cat [F. libyca] is
a competitor of the fox. Death of red foxes due to encounters with
carnivores and competitors constitutes 2—12% of the total number of
deaths (more often in forest and desert zones).

paz

Table 44. The ratio of species-specific to generalist helminth species in the
foxes in the Crimea (from Kadenatsii, 1957)

In common with other species

Total Species-
helminth specific With With With With With
species dogs cats badger steppe beech
polecat marten
34 450 29 12 8 None 2
Wied 88.5 95:2 2BES 5.9

Population dynamics. Numbers of the red fox are not stable
(Formozov, 1935; Chirkova, 1941, 1947, 1947a, 1952a, 1953, 1954;
Teplov, 1949, 1960; Shcherbina, 1961). In favorable years, minimal
density of the fox population occurs in spring after the end of the
harvest. A maximum number is attained with the conclusion of repro-
duction and is usually observed in autumn after the emergence of the
fox cubs from the burrow. Rarely is the increase in the population
more than 3—4 fold (taking an average in such years of 7 to 8 cubs
in a litter). In unfavorable years, especially with an epizootic out-
break, the foxes which remain until spring, with worsening conditions,
produce a few weak offspring which soon die. In this case, the popu-
lation in autumn may display even fewer numbers than in spring.

The multiannual numerical dynamics of the red fox are reflected
by the well-known data on tanned skins. Tanned skins of red fox has
changed within three-fold limits from the middle of the 20’s to the
50’s of the twentieth c[entury]. The rise in number of foxes took
place in the mid-20’s and 30’s. The minimum number and weak
harvest arrived at the beginning of the 30’s. Unusual abundance of
foxes was observed in time of war—in the mid 40’s, when, beginning
with 1942/43, tanning quickly grew until 1946/47, after which it
decreased. A new increase was noted at the beginning of the 50’s.
During the post-war period, tanning of red foxes increased by 150%
as compared with the prewar years. The increase in tanning was
promoted not only by the general growth of the fox population in all
natural zones of the country, but also by intensification of harvest
(Fig. 140).

In the numerator—number of helminth species; in the denominator—percentage
of generalist helminth species of a given domestic form.

375

553

Near the northern limits of the range, fox populations vary quite
sharply every 2—4 years, following the fluctuations in numbers of

-small rodents. In the Nenetsk National Region, tanning of fox skin

usually grows in the course of 1—3 years and sharply decreases in one
year; often, they co-decline with the dynamics in Arctic fox numbers.
On the Kola Peninsula, the numbers of red foxes depend upon mass
reproduction of red-backed voles, which is repeated nearly every 4
years (Koshkina, 1957; A.F. Chirkova).

In the taiga zone in the northeastern European territory of the
Union, in the Pri-Yenisei taiga and in Yakutiya, red fox numbers
depend on reproduction of arctic hares (Middendorf, 1853; Teplov,
1949; S. Naumov and Popov, 1957). In Yakutiya, the fox populations
quickly increase in years of “outbreak” of this rodent*, in the 2nd—
4th year after its mass reproduction. The increase is repeated after
8—12 years, not synchronously in various regions of Yakutiya (Labutin,
1960). In several regions of the north, there is a significant abundance
of forest game birds. In the forest zone, the peak in fox numbers
occurs in years of decrease in number of hares and game birds,
coinciding with reproduction of small rodents. Under such conditions,
for example in 1943/44, tanning of fox skins in Yakutiya increased
by 310%, attaining in 1944/45 22.8 thousand items, while the usual
increase did not exceed 200% and the average number of tanned
[skins] per year (for 34 years) consisted of 5.8 thousand items. In the
Nizhne-Kolymsk region, from 20 to 700 skins of fox were tanned in
various years; the annual catch of foxes increased sometimes by 3—
4 times. In Pechoro-Ilychsk preserve, fox numbers changed 8—12 fold
in the course of several years (Teplov, 1949). In Tatar Republic, 5
increases in fox numbers were observed in the last 40 years, at inter-
vals of 7 to 9 years (V.A. Popov). In the Far East, growth of fox
numbers is connected with the migration of spawning salmon into the
rivers, which usually occurs biennially, as well as on the abundance
of small rodents; other kinds of food are also significant (A.F.
Chirkova).

In the forest-steppe and steppe zone, the dependence of fox
numbers upon small rodents, mainly the common [M. arvalis] and
social [M. socialis] voles and sagebrush voles [Lagurus] is precisely
evident. Tanning of red fox skins in successive years increased by
300 to 400% in the Altai territory and Penzensk district, by more than

*Sic; hares are no longer considered rodents, but lagomorphs—Sci. Ed.

376

375

554

400% in Saratov district and Stavropol’ territory, and by 520% in
Rostov district. During the extent of the postwar years, tanning varied
in Voronezh and Volgograd districts 6-fold and in Rostov district—
10-fold. In 1956/57, as a result of food shortages, tanning of red foxes
in Orenburg territory decreased 10 times. On experimental areas in
Kiev and Voronezh districts and in Stavropol’ territory, the number
of foxes changed 3—4 times in two successive years.

In the southern deserts of Kazakhstan and in Middle Asia, red fox
numbers changes during a 24-year period, depending mainly upon
reproduction of several species of gerbils and voles. Peaks of
skin tanning are repeated here about every decade. The amplitude of

Thousands
iment of skins

Number
of foxes

1937 1938 1939 1940
193 я 3
755/- в 9399 33 940,

1936

Fig. 139. Counts of red fox in an experimental plot, prognosis of numbers and

general catch (tanned skins) around Stavropol’ territory, Cis-Caucasian steppes

(from Chirkova, 1947). 1—Number of foxes by count of litters in spring; 2—

Fox catch in experimental plot; 3—Tanned skin in the territory; 4—Prognosis

of numbers in the direction of forecast decrease or increase of population.

Left—number of established individuals according to No. 1. Right—thousand
skins.

555

fluctuations in tanning in Turkmeniya and the Uzbek SSR changed
not more than 2 times during the postwar years. In montane regions
of the south, red fox numbers change depending on the reproduction
of montane voles. In Armenia, the Tien Shan and the Pamir, fluctua-
tions in fox numbers is frequent but within a small range, which
corresponds to variation in populations of mouse-like rodent. In
Tadzhikistan and Kirgiziya, tanning in the postwar years fluctuated 2
times; in Dagestan, 2—3; and in Armenia, 3—3.5 times (А.Е. Chirkova).

The influence of harvest on the dynamics of fox numbers has
been little studied. This effect depends on the number of hunters, the
time of harvest and the instruments of capture employed. On an
experimental area in the former Spitsevsk region of Stavropol’
territory, hunters each year caught from 24 to 100% (average, 78%)
of the counted fox population. In Brovarsk region of Kiev district, on
a permanent plot of 400 km’, 53.5% of the foxes counted in summer
were caught in the winter of 1951/52. Catch of foxes within this limit
does not, apparently, undermine the main parent population. Only in
1954 and 1955, after a heavy winter in Stavropol’, was there ob-
served a considerable decrease in the fox population, harvest sharply
decreased, and starting from the middle of winter, capture of fox was
prohibited in 12 regions of the territory; after this the population of
animals was again quickly restored. An analogous case of
overharvesting took place is Yakutiya in the beginning of the 50’s;

1924 и? Г

92991931 1935 1935/6 1945 /yg 1955/56

Fig. 140. Fluctuation in tanned skins of red fox within the whole Soviet Union
as an indicator of the fluctuation in numbers of the animal (A.F. Chirkova).

1960/5)

556

after a temporary prohibition on capture, in the 3rd year tanning again
regained its usual level (A.F. Chirkova).

The abundance of the red fox, which was evidenced everywhere
in postwar times, is explained both by mass reproduction of small
rodents, and also by the absence of regular harvesting.

Field characteristics. Tracks of red foxes are characteristic and
easily distinguished from those of related species. The gait of foxes
is direct (straight as a string) alternating footprints. In an easy trot or
“trusts [jog-trot]” by which the red fox most often moves, it places
its anterior left foot and the posterior right exactly in one place, and
then, stepping forward with the right anterior one, it lowers in its
place the left posterior foot. In this way the impression is given that
a bipedal animal has passed by. The distance between footprints is 20
to 30 cm. With a slower gait, when the fox sneaks up on its prey,
looks out for danger, or satiated, leaves its food for a rest, the dis-
tance between tracks decreases until they nearly contact and the track
of the hind foot is slightly behind that of the forefoot. When excited
or disturbed, the red fox proceeds in large bounds. In deep snow, it
moves in a short gallop, in which the tracks of all four feet remain.
Running from danger, it careers in an extended gallop. In this case,
it can make a jump of up to 4.3 m (Dinnik, 1914) (Fig. 143).

The footprint of the red fox is similar to that of the dog, but more
compressed, narrow and, not as wide-pawed; the contours of the
lateral digits do not touch the posterior margin of the middle digits
and do not cover them from the sides as in dogs. On a thin layer of
soft snow or on wet soil, the prints of the claws are always visible.
Measurements of the elongated track of the foot are 4—5 x 6-7 cm
with claws (Fig. 142). Loading on 1 cm? of the track surface is 40 g
or more in the middle belt [of the Union] and 28 g in the Pechora
[region] (Formozov, 1959; Teplov, 1960).

The female track is smaller, narrower and more pointed and the
stride is shorter; in young, the tracks are still smaller and not so well
marked as in females. Anterior drag marks and posterior drag-marks
on the snow made by the shorter-legged female are, naturally, more
frequent and longer. The female often proceeds by hiding behind
bushes and irregular relief while the male usually walks directly
forward, not hiding. In time of rut, the female leads the males. The
sex of foxes can be determined by the manner of urination; with
males usually urinating near protruding objects—rocks, hummocks or
bushes.

557

Fig. 141. Track of red fox crossing a marsh in snow of moderate depth with
heavy frost. Oka preserve, Ryazan district. January 1950. Photograph by
V.G. Heptner.

According to hunters of East Siberia, tracks of melanistic indi-
viduals, especially blackish-brown foxes, by comparison with red are
differentiated by their larger dimensions, not only due to the larger
size of the feet, but also by their greater fluffiness. Melanistic foxes
are more cautious than red (Slyunin, 1900; Buturlin, 1913). Hunters
consider that the better furred the fox, the more frequent are its
tracks. Individual differences in tracks exist, for example, “Japistost’*

*An uncommonly used trapper’s term—Sci. Ed.

378

558

when the heavily furred pads of the feet leave a wide and diffused
track.

The direction of the paths of a fox on the deep or drifted snow
can be determined by the drag marks. The latter, produced when the
foot is raised from the track is usually shorter and more abruptly
lifted from the snow, breaking its edge; snow pellets are thrown
forward. The drag mark formed while setting the foot onto the snow
is elongated and sloping.

A series of fox tracks extends in a patterned manner, traversing
a field, meadow, or shrubland, passing through hills, descending into
a ravine, following the edge of a wood, sometimes along ski-tracks,
often with stops along the way. During times of food-seeking, the fox
walks in a’ zigzag manner, moving first to one, then the other side,
examining each hummock, clump of grass, and bush met with, walk-
ing around haystacks and dung-heaps, and not neglecting a single
object on the snow surface in its way, sometimes returning along its
own tracks. The red fox does not make trails, since on passing for a
second time or repeatedly along its tracks, it places its feet accurately
in the place of the previous tracks. Trails are made only by young
foxes, usually along banks of small creeks overgrown with reeds. At
different times of day, the red fox traverses particular paths and on
bright days it hides more often than on cloudy ones.

In years with abundant small rodents, on the path of the well-fed
fox, holes are left in the snow through which it hunted voles. In deep
snow, these “kopka” [diggings] are up to 80 cm deep (Lapland pre-
serve; Nasimovich, 1948). On a 2-km path up to 30 holes were found
(Formozov, 1959). Along the tracks of foxes, killed and mauled shrews
and even moles, which it usually does not eat, are often found. Shells
of eggs, sucked in summer and rotten in winter, have two holes made
by the canines of the fox at a distance of 1.5 cm from each other. The
remains of a large or moderate-size bird are usually encountered in
the form of the whole wings or separate remiges or tail feathers, with
the quills severed at the base. The red fox, upon capturing a hare,
usually does not completely eat it, but digging a hole in the ground
or snow, puts the remaining part of the carcass there and treads and
kneads the surface with feet and nose.

The lair of the fox is found sometimes under a bush, or near a
hummock, often on its summit, on a haystack or thatch pile, and even
on a bent tree at a height of some meters from the ground. In forested
localities, the red fox prefers for resting coniferous forests—spruce

379

378

559

forest, where it can hide among the shaggy boughs, pine forests with
mossy bogs where it can hide behind hillocks. From one lair to
another, tracks of the fox may extend for some kilometers. Some-
times, before retiring to rest, the fox executes a trick—a sideways
jump into any sort of shelter. The dimensions of the lair in the Lapland
preserve are 40 x 50 cm in width and 10 to 15 cm in depth
(Nasimovich, 1948).

Except at the breeding period, red foxes rarely frequent burrows;
only in extreme cases, in times of wet weather, during severe frosts,
or when hiding from danger, does the fox enter a burrow.

In the second half of winter, paired tracks of foxes are often
encountered, and at time of estrus, straight away several animals are
together. With this, one may observe the signs of fights among males,
and remains on the snow of tufts of hair, even clots of blood. In
spring, sticky secretions may be found on the tracks, characteristic of
estrus, and in years with food deprivation and at the beginning of
epidemics, bloody mucus may be found.

Feces are usually 1.5—2 cm thick and 3-9 cm long, closely cov-
ered with a film of mucus and well-formed, sometimes with a
constriction; in fresh form dark-olive in color, and on drying—whit-
ish and earthy. More often, it consists of fur of rodents, sometimes
with a mixture of small feathers, large hairs of domestic animals,
pieces of bones, parts of insect chitin, fragments of dry plants and

A К

Fig. 142. Track of red fox on clay soil. Manych. 28 May 1940. Sketch by A.F.
Chirkova, natural size.

560

others. In summer, especially with small numbers of rodents, feces
are met with that are entirely composed of chitinous parts of insects,
which fall apart in the hand. In the floodplains of the Volga and
Kama, with insufficient animal food, foxes in winter eat rose-hips
and their excrement then has a brick-red color (V.A. Popov). Near
the den burrow, at points around the main entrance, is scattered
excrement of the fox cubs, 8 to 15 mm thick, depending upon age and
sex. Here also occur the discarded remains of prey brought by the
[parent] foxes. Narrow paths lead from the den burrow, which the fox

ny we
dy lao ae

&)
о

(Y:)
Ly
[С в №
®

a2 =>
3

Src ogee и ЕС

=
о See
St
=

bt dae
i whe №
! 2 3 4

Fig. 143. Different gaits of the red fox (from Когпееу, 1956): 1—Walking, light
trot; 2—extended trot; 3—trot with “paired limbs” [pacing ?]; 4—gallop.

380

561

cubs trample out on various sides. On these as well, one may find
pieces of feces and remains of the fox cubs’ food.

The red fox orients itself by sound; it can hear the sound of
blackcock changing roosts at about 600 paces, the flight of a crow—
at 1/4 to 1/2 km and the squeak of a mouse—at around 100 meters
(Zvorykin, 1931). But cases of completely deaf foxes are known in
nature (Postnikov, 1936). Sight in the fox is well developed, but it
reacts mainly to moving objects. Scent is good, but weaker than in
specialized dogs. Apparently, olfaction operates at close range. The
fox shifts objects along the ground not only by its feet but also its
muzzle.

The voice of the red fox may be heard before storms, in inclem-
ent weather, and severe frosts, but most often at the beginning of the
breeding period, when the female calls males by barking; usually at
the beginning of twilight and until morning (p. 532). After
2-3 dull, metallic barks resembling the voice of small mongrel, comes
a short montonous howl. The female barks while sitting, raising its
muzzle upwards. Males do not respond immediately: their barking is
more frequent and impatient and is interrupted not by howling, but by
a yelping sound resembling the cry of a jay or fighting cats (A. Ch.).

Practical Significance*'

The red fox is a valuable fur-bearing animal. In the past century,
more than one hundred thousand foxes were caught annually in Russia.
In the beginning of the twentieth century (1907—1909), according to
data of the International Fur Trade, the yield from fur bazaars in
Russia was 363 thousand red fox skins including sivodushka,
chernoburaya and karaganka foxes.

In the Soviet Union, from 1924/25 to 1958/59, an average of
487.3 thousand red foxes were caught yearly. During the post-war
years, tanning considerably increased. During a 35-year period, about
17 million skins were prepared. In fur manufacture, red fox occupies
fourth place. The greatest number of skins tanned are from steppe
and desert regions; at the same time, the taiga zone yields more costly
fox fur of better quality. One hunter in the forest zone frequently
catches only occasional animals, rarely more than ten, while in the

'Silant’ev, 1898; Brass, 1925; Brauner, 1930; Chirkova, 1941, 1948, 1963:
Korneev, 1956; Ling, 1956.

—

562

steppes and deserts, he gets several tens and in some years, even
hundreds of foxes.

The fox destroys harmful rodents—voles, mice, ground squirrels,
hamsters and others, which has especially great importance in agri-
cultural regions. Eating about 300 g of meat per day, each fox can
destroy about fifteen voles” and in one year not less than 3000 voles
and mices, or 1000 gerbils. During the three summer months, one red
fox together with its litter can destroy more than 500 ground squir-
rels. It destroys also poisonous snakes. Among insects sometimes
eaten by the fox when abundant, especially in steppes, deserts and
montane regions of the south, the majority are locusts, termites, cock-
chafer beetles, weevils, darkling [tenebrionid] beetles, larvae of beetles
and moths which belong to the pests of agriculture, fruit orchards and
forestry. In southern Turkmeniya, analysis of animal food of the fox
characterizes it as an unquestionably useful animal (Table 45).

In the period of low numbers of small rodents, the fox destroys
many more than in times of abundance. Therefore the value of fox
as a destroyer of agriculture pests is especially great in the years of
their depression, since it hinders the quick restoration of their num-
bers.

Accused as a predator, the fox is implicated in the destruction of
game and song birds, hares, muskrats and the young of ungulates.
The harm caused by this predator is especially appreciable in pre-
serves, reserves, and hunting farms where game birds such as
pheasants, partridges, grouse and others, which are ground nesters,
are protected and raised, as well as in the neighborhood of poultry
farms. However, often the charge of game destruction is exaggerated,
since the degree of fox damage is usually determined by the remains
of food found at its burrow. The limb bones of hares or the wings of
large birds may be preserved for a long time leading the observer into
error. Thus, in North America, where rabbits (Sy/vilagus) smaller
than our species of hares are often eaten by red foxes, the remains
of the comparatively large animals (piglets, hares, pheasants and do-
тезис chickens) were found near burrows 11 times more often than
in the excrement, while the remains of the meadow vole [М.
pennsylvanicus| were found 75 times less often (Errington, 1935).
It is also known that single individuals or separate litters of fox

In the stomachs of fox, up to 20-21 voles which it feasted on in a nest, were
found (Formozov and Osmolovskaya, 1963).

563

specialize in eating several species of secondary importance, as for
instance, domestic fowl (Pavlov et al., 1961; Ishunin, 1962).

The fox captures relatively few brown hares [L. europaeus] and
destroys arctic [Г. arcticus] and tolai [L. tolai] hares in considerable
numbers only in years of massive diseases and plagues of these
lagomorphs*. In Yakutiya where hares are characterized by sudden
increase in numbers in the course of 2-3 years and also sharp de-
creases, these carnivores are not the cause of the periodic decrease
in the number of hares, since its population in years of abundance of
these lagomorphs* is insignificant. When signs of depression appear
in hare populations, and epizootics develop, fox feed freely on them
(more than 50%, and up to 96% of occurrences in the food of indi-
vidual litters), and so do the majority of predatory animals and birds.
Under these circumstances, the red fox serves as a factor of great
importance in the sense that it rapidly removes carcasses of dead
hares. Later, in the period of minimal numbers of hares, the fox as
well as other predators, holds back a new wave in the growth of
population of these logomorphs* (Labutin, 1960). In some regions,
the red fox kills many young ungulates, for example roe deer in
preserves in Crimean region, but it destroys mainly weakened young,
thus itself increasing the health of the population. The role of the red
fox in destruction of small song birds is not great.

On the negative side, there is the role of fox in the transmission
of several dangerous infections, especially rabies as well as carnivore
distemper, wild rabies in the north, and several other diseases. The
significance of the species in this respect is still not sufficiently
studied. The fox is also dangerous as a vector of several species of
helminths—alveolar echinococcus, trichina, tapeworms and others, as
well as ectoparasites—mange mite, fleas and ixodoid mites—carriers
of blood parasites and other infectious diseases of humans and
domestic animals.

Techniques of catching the red fox are of various forms (Silant’ev,
1898; Sabaneev, 1904; Zvorykin, 1935; Chirkova, 1945; Gerasimov,
1950 and others), but basically in the greater part of the Union, it is
trapped and shot. Methods and manner of setting traps in fox trails,
in the places where it catches its food, in hare trails, and in deep
snow—under the track; under ski or sled tracks where the fox is not
afraid to walk along them—are exceedingly variable. Traps are also

*In Russian original, “rodents’—Sci. Ed.

564

Table 45. Correlation of vertebrate and invertebrate animals having different
economic significance in the food of Badkhyz foxes (according to multiyear
data; Shcherbina, 1961а)

Economic Vertebrates Invertebrates
importance Absolute % Absolute %
number number
Useful 799 Dilies aii 11.6
Neutral 12 0.3 4 0.06
Harmful 2687 Wey 5549 88.3
Unknown economic
importance 253 6.7 —— —
Total 3751 100.0 6284 100.0

set making use of peculiarities of relief and landscape—near rocks,
lone standing trees, bushes, etc., on sites of old campfires, and near
frozen sections of creeks and small rivers, where the fox crosses
them. When a fox as determined by its track is found in a burrow,
the trap is set at the hole. Traps are also set out near baits, in particu-
lar odiferous ones; sometimes live baits in the form of a small bird
Or mouse are used with them. A better method of setting traps is at
lures*. On the tributaries of the Ob’ [river], traps are placed near
unfrozen parts of the river. In the south, traps are situated in former
livestock corrals, and in the sand of southern Kazakhstan and Middle
Аз1а— ипдег bushes and under the nests of raptors.

Red foxes are ambushed at lures, by preparing in advance a place
for sitting or a dug-out. They are hunted with a decoy mouse squeak
or the cry of an injured hare. Hunting fox by beaters with flags is
widespread. At the time of estrus, males are driven out and shot. In
the forest zone, stalking is employed. In the south, they are hunted
with guns along chernotrop** among the furrows of ploughed fields,
where foxes like to mouse, and also in drives, the foxes being forced
out from reed thickets of the steppe rivers. Hunting with hounds is
widespread; it is only possible before deep snowfall, and is successful
in places with varied landscape. Foxes are hunted by setting fox-
terriers and dachshunds on them in burrows. A specific kind of fox

*The distinction between the Russian words “primanka—bait,” and ‘‘privad—
lure,” is not clear in this context—Sci. Ed.
**[ iterally, “black path’; refers to bare ground in winter—Sci. Ed.

383

565

hunt is found in the steppes and semideserts of Kazakhstan and Middle
Asia, with golden eagles and borzoi (“1а2у”). In the Trans-Baikal
steppes, foxes are hunted by driving them with horses.

In the greater part of the territory of the Soviet Union, the red
fox, as well as other commercial species, is protected. Protective
measures for the fox mainly include timing of the hunt, which is
permitted only in winter. Harvest usually begins in November and
continues to the end of February. Hunting seasons change depending
upon the latitude of locality, and annually depends for its timing on
the maturity of the pelage.

In areas where poultry breeding and sport hunting of forest game
birds and arctic hares is developed, and also where foxes are severely
infected with mange mites, the animals are killed during the course
of the entire year, for instance in Estonia; from 1958, shooting of
foxes was permitted year-round in the Ukraine and Georgia (which
was not justified). Foxes were killed in muskrat farms also.

Organized effective shooting and trapping of foxes throughout
the year using various methods (except those generally dangerous)
must be permitted only in individual localities or in individual years
when this species carries an obvious danger to the economy, or be-
comes dangerous in connection with the possibility of spreading
infectious diseases. In other cases, it is necessary to promote the
increase of the fox pack. Only a planned economy allows the possi-
bility of avoiding extreme decision in respect to the species as a
whole, to use it in the best manner. In Bulgaria and several countries
of central Europe, the red fox is considered a useful species in agri-
culture and forestry. In a series of countries—Australia, Bulgaria,
some states of the USA, Canada, Romania, Finland, and Yugoslavia,
the taking of foxes is not limited, and in some states or estates a
bounty is paid for their destruction. At the same time, according to
some studies carried out in Europe and in the USA, it is acknowl-
edged that unlimited extermination of the red fox and other predators
does not confer a benefit on the protected species which serve as their
natural prey, since these species begin to over-reproduce.

Acclimatization measures with red foxes were begun in Russia in
the 17th c[entury], when the black-brown foxes were brought from
Siberia tc forests in the vicinity of Moscow (G. Dement’ev, 1956). In
the 30’s of this century, the silver-black foxes from farms were re-
peatedly released into different regions of the Soviet Union—Northern
Territory, Lithuania and others (Parovshchikov, 1936; Ivanauskas,

566

1956 and others). Naturally, this benefited nothing. The transfer of
the common red fox to other continents took place in the 17th and
18th c[enturies]: red foxes were sent from England to the U.S.A.
Foxes brought into Australia for the control of wild rabbits acclima-
tized well, and resulted in its coat becoming somewhat modified. But
the result of acclimatization was shown to be negative, in that the fox
began to destroy not only endemic birds, but also caused great harm
to populations of small kangaroo (Zhitkov, 1934, 1940; Presnall, 1958;
Kastelyarnau, 1960*; Nasimovich, 1961).

The keeping of foxes in captivity was practised in previous years
in the regions of the extreme North and in some districts of Siberia.
Young foxes were taken from burrows and kept in primitive log
coops or restrained until the time of killing. Due to the low value of
skins thus obtained, this was prohibited. Commercial fur farming in
the USSR has existed since 1928 and has been greatly expanded in
the postwar years. American silver-black fox and forms derived from
them (platinum, white-faced and snowy) are bred in captivity.

Commercial fox-raising abroad, especially in North America and
in Scandinavian countries of Europe began even at the end of the past
century, and by the first half of the 20th c[entury], it was very widely
developed. Since the time of the Great Patriotic [2nd World] War, the
breeding of foxes has considerably diminished as a result of the fall
from fashion of long-haired furs, but in recent years, it is again being
restored (A. Ch.).

Subfamily of Red Wolves
Subfamily Simocyoninae Zittel, 1893
Genus of Red Wolves
Genus Cuon Hodgson, 1838

1838. Cuon. Hodgson. Ann. Mag. Nat. Hist., [.р. 192. Canis primaevus
Hodgs. = C. alpinus Pall.

1839. Chrysaeus. H. Smith. Jardine’s Nat. Libr. Mamm., 25,
р. 167. Canis dukhunensis Sykes = С. alpinus Pall.

1888. Cyon. Blanford. Fauna Brit. India Mamm., I.p. 142. Correction
of Cuon.

*Not in Literature Cited—Sci. Ed.

384

385

567

1888. Anurocyon. Heude. Mem. Hist. Nat. Emp. Chin., 2. p. 102.
Anurocyon clamitans Heude = C. lepturus Heude = C. alpinus
Pall. (V.H.).

Forms of large dimensions.

Skull heavy and massive, relatively short with broad, widely
separated zygomatic arches, high (height of occipital region almost
equal to half condylobasal length of skull); in adults, protuberances
and crests well developed including sagittal crest.

Facial part of skull greatly shortened, massive, relatively wide in
proximal part—distance from anterior margin of premaxilla to preor-
bital foramen is less than skull width at level of molars. Interorbital
(frontal) region convex, this convexity (related to formation of air
sinuses) extends forwards to basal half of facial part, resulting in a
convexity instead of a depression on profile line at level of anterior
margin of orbit. Therefore, “frontal process” is absent and whole
upper profile of skull represented by quite flat convex arch. Several
depressions occur only in middle part of nasal bones.

Posterior margin of nasal bone extends far behind posterior bor-
der of frontal processes of maxillary bone. (In other representatives
of the family they lie at approximately the same level or the opposite
relationship is observed.) Inner parts of anterior ends of nasal bones
form blunt projection in middle of nasal notch. Supraorbital processes
relatively small with weakly defined depression on upper surface;
their ends are bent downwards. Posterior margin of toothrow extends
to level of anterior margin of interpterygoid depression. Deep notch
and protuberance on lower margin of lower jaw in front of angular
process absent.

Dental formula 12clp4m2 — 40. Teeth quite strong, but first
За 2

upper molar is weak (approximately two times smaller than in wolf),
its inner blade small and carries only one cusp (in other members of
the family, they are 2 to 4). On first lower molar (carnassial tooth),
only one small cusp behind tall main cusp.

Canines quite massive, slightly curved and short—when jaws are
closed, ends of upper [canines] do not reach or only just reach middle
of mandibular bone; ends of lower [canines] reach margin of alveolus
of upper [canines]. Tips of upper canines directed straight downward;
the sharp cutting edge on posterior surface of canine and ribbing on
lateral surfaces absent. Cutting edge of incisors without accessory
lobes.

568

Limbs moderately long, trunk proportional, not excessively elon-
gated; tail quite long, extending below tarsal joint, but in standing
animal, does not reach ground. Pads of digits III and IV united to the
rear, and form horseshoe-shaped figure open in front'. Color uniform,
quite bright, reddish.

“at
i

Ei >

3 X : 4 Moscow
[о

Fig. 144. Range (reconstructed) of genus of red wolves, Сиоп Hodgs., and species
range of the red wolf, Cuon alpinus Pall. (V.G. Heptner).

'This character, noted in material of Zoological Museum of Moscow University
must be visible in the track (Fig. 17, p. 76).

“AOIeWOY “N'Y Aq SuNuIeg “(dss иошшр 50) тецуде Summsind “[OZ 3 ‘ивуу пма@5ац пиар uoND ‘sarjom pay “¢ эаа

386

Fig. 145. Red wolf, Cuon alpinus Pall. in winter fur. Specimen from China in
Moscow Zoo. Photograph by A.P. Zhandarmov.

There are 6 or 7 pairs of teats, occasionally 8. Sexual dimorphism
is absent; seasonal dimorphism clearly demonstrated in northern forms,
in southern, weakly or not manifested.

A forest, montane forest, and in part montane form. In vertical
direction they attain extreme heights. Young number 2—4~7, in ex-
ceptional situations, more. Young are born in burrows, clefts in rocks,
and similar places, sometimes forming even “colonial” settlements.
Predators, feeding on flesh, mainly of ungulates. Even in the breeding
season they remain together and hunt in packs, sometimes quite large,
bringing to bay their prey.

The range of the genus is quite considerable and occupies the
central and southern parts of East Asia. In the north, the range ex-
tends to the Amur region, Trans-Baikaliya, Altai, Tien Shan and
Pamirs. In the south, it occupies Hindustan, Indochina with the Ma-
laccas, Sumatra and Java (for more, see beyond in species description).

570

In its systematic characteristics, the genus is quite isolated from
the other genera inhabiting the USSR, which is emphasized by plac-
ing it in its own subfamily Simocyoninae. Even if the division of the
family Canidae into subfamilies is not accepted, the isolation of the
genus from the above-considered assemblage must be regarded as
significant. The general appearance of the animal is also character-
istic (see below). It is usually accepted that the red wolves stand
closest of all to the genus Lycaon (African hunting dogs), although
the latter are markedly differentiated by dental formula (the same as
in Canis), the color, and a series of other characters.

Reliable fossils of representatives of this genus are known from
the Pleistocene of Asia and Europe. It is undoubted, however, the
genus diverged and was established much earlier, and already
existed in the Pliocene. There are indications of Pliocene finds in
Italy. In any event, representatives of the subfamily are known in
Europe from the upper Oligocene, and are near to the genus Cuon in
the lower Pliocene. In the Pleistocene, and in part in the Holocene,
red wolves were more widely distributed in the north—they lived also
in the southern Urals, in the Ukraine, and westward to Czechoslova-
kia, Switzerland, Italy, France, Monaco and Spain, while eastward,
they were found in the Holocene at Nizhne Udinsk.

The generic independence of the group is without doubt and is
accepted by all investigators. The recent suggestion (Haltenorth, 1959)
to unite the genus Cuon with the genus Lycaon cannot be accepted
even in the very widest interpretation of the genus.

Until recent times, several species were included in the genus,
but now it is accepted that it includes only one species with several
races. This constitutes about 3.4% of the species in the family.

In the USSR fauna there is one species, the red wolf Cuon alpinus
Pallas, 1811, 1.е. about 0.3% of the species of mammals in the USSR.

The range of the genus in the USSR occupies the Amur-Ussuri
territory, the montane southern fringe of Siberia, and the mountains
of the eastern part of Middle Asia (V.H.).

571
RED WOLF*
Cuon alpinus Pallas, 1811

1811. Canis alpinus Pallas. Zoographia Rosso-Asiatica, 1, p. 34.
Udskoi stockade.

1935. Cyon alpinus hesperius. Afanasjev et Zolotarev. Izv. Akad.
nauk SSSR, otd. matem, 1 est. nauk., No. 3, p. 427. Aksaisk
Plateau, Tien Shan.

1936. Cuon javanicus jason. Pocock. Proc. Zool. Soc. London, p. 51,
Altai.

Diagnosis
The only species in the genus.
Description

In general appearance the red wolf is unique and is not similar to the
gray wolf—ain its constitution, rather, mixes features of the wolf (or
jackal) and fox. It is an animal with a somewhat elongated, but
proportional trunk. The tail is long, nearly half the length of the trunk,
387 and in full winter pelage, almost reaches the ground’. It is covered
with very long, dense hairs which appear very thick (thicker than in
the red fox).
The winter fur is dense, and quite soft and very long. The hairs
on the back are especially long, where individual [hairs] reach 160
mm. On the cheeks and along the upper neck the hairs are
elongated. Therefore, the head in winter fur looks small. It is wide in
the zygomas, the muzzle is quite short, but pointed. Ears are large
with blunt, even somewhat rounded, tips, protruding well out from
the fur. Their bases are strongly approximate and they are, therefore,
situated high on the head. All of this gives the animal’s head
a completely unique appearance, and expression which is similar
neither to the red fox nor to the wolf.

*In English, also dhole—Sci. Ed.

п the red wolf (in captivity) there is a completely distinctive manner—the
whole appearance expresses timidity and tension, the animal seems to be flattening
itself and to be slightly lame in the hind legs. With this, the end of the tail drags on
the ground [see Fig. 145].

The general color tone of the whole fur is reddish, noticeably
variable individually, and apparently, in part geographically (within
the USSR). In the animals with brightest color in winter the back is
clothed with saturated rusty red-to-reddish color, brownish highlights
occur along the top of the head, neck and shoulders; throat, chest,
sides, belly and upper parts of the limbs are less bright, and more
yellow in tone. The lower parts of the limbs are lighter, yellowish-
whitish, and on the anterior sides of the forelimbs dark brownish
bands are weakly marked. The muzzle, area between the eyes, and
the forehead are grayish-reddish, with color brighter on the forehead.
The extraordinarily luxuriant and fluffy tail has a rusty-brownish
color and is covered with brownish highlights formed by the dark tips
of the guard hairs. At the end, this highlight is denser and the tail
darker. The underfur and the bases of the hairs are in color gray of
various intensities, sometimes with a more or less marked touch of
yellowish.

Paler and duller colored individuals have a tone from ocherous
reddish to grayish-rusty. Chin, throat, bottom.and sides of neck, belly
and the lower parts of the sides are white with a more or less devel-
oped ocherous or light ocherous highlight. The tail is reddish-ocherous
in its main part, darker due to the brown tips of the hairs in its distal
half and especially at its tip; below it is lighter. In particularly light-
colored individuals, darkening of the tail is almost undeveloped.

In some individuals, the general color is dull with weakly devel-
oped rusty-red tone.

Summer fur coat is much shorter than the winter, coarse
and darker, brownish. The tail, however, is covered with fluffy long
hairs.

For characteristics of the skull, see description of the genus.

Body length is 103 cm, tail length 48 cm. Hind foot length
22 cm, ear length 8 cm’.

Greatest length of skull 171.0-200.0 mm; condylobasal length of
skull 174—188 mm; zygomatic width 103—118 mm; width of cranium
61.0—68.3 mm; skull height in region of tympanic bullae 70—73 mm,
length of muzzle from orbit to the alveolous of incisor 70.0—81.5 mm;

’Measurements of one specimen from Ussuri territory (Ognev, 1931). For red
wolf from the Himalayas С. a. primaevus, body length is given as 92—96 cm and
weight, 17.3 kg (Pocock, 1941).

575

SS) a
BS Sy
= SSS
5 SS

7/7
,
Z2

Md,

Weegee

ИГ

aa =
ONT Ass
117

it
SN Wes

Si
XSAN

<
: QD

Lb, Pp
4,

>
<

Sys vy
Ny
if

Ass 0 yi
<<) y

= —>S
WSS vty = GY.
Ne WN) NS Я LG
м У У И А SAN % ay
: UNOS
5 |

2
7
< (г.

ann <
AY) SS =
Ах
А К
== as \a rs и:
И = sear!

хх

388 Fig. 146. Skull of red wolf, Cuon alpinus Pall.

389

574

length of nasal bone 53.2—71.5 mm; length of upper tooth row 74.0—
80 mm.’

In connection with the rarity of the red wolf in the great part of
its range in the USSR, the number of skins and skulls in our museums
is very few. Therefore, it is impossible to give a sufficiently full
picture of variation in this species in our country (V.H.).

Systematic Position
Only species in the genus.
Geographic Distribution
South, Central, and southern half of East Asia.
Geographic Range in the Soviet Union

This represents the narrow northern margin of the range and occupies
a negligible part of the country—the southern fringe of the Far East,
eastern and middle Siberia, and the eastern part of Middle Asia.

In the east, the northern border of the range of the red wolf
begins at the shore of the Pacific Ocean in the region of the lower
Uda (Udskoi stockade).

In spite of the assertions of some earlier authors (Shrenk, 1858;
A. Nikol’skii, 1889) perpetuated also in recent literature (Ognev,
1935; Ellerman and Morrison-Scott, 1951), the red wolf is not
and never was on Sakhalin. From Uda, the range boundary goes
westwards somewhat to the south of the Stanovoi range and passes
a bit to the north of the northern bend of the Amur, not reaching the
main axis of the Stanovoi range here (in Gilyui and Ol’doi, this
species is not recorded®). This is the most northerly habitat of the red

‘Figures according to somewhat contradictory data in literature (Ognev, 1931;
Afanas’ev and Zolotarev, 1935; Novikov, 1956) and they may give only a general
representation of skull dimensions.

‘Information concerning the penetration of the red wolf into the Stanovoi moun-
tain range given from old authors (Maak, 1861; Radde, 1862) is somewhat contradictory.
Apparently, it should be understood thusly: to the west of the range in the region of
the northern bend of the Amur, the animal penetrated nearer to the main axis of the
range than in the east, where it apparently did not spread farther than the foothills
(according to Radde, it was also rare in the Bureinsk mountains). It must be taken into
consideration that the known contradictions of the proposed text with the texts of the
referenced authors and that of Ognev (1935) are explained not only by some new data
and corrections, but also terminology—not too long ago, the Stanovoi range was
frequently called Yablonovoi. This appears as an error of some contemporary authors.

391

575

wolf in our country and the most northern part of the range of the
species.

Extending farther, to the north of southeastern Trans-Baikaliya
and the upper Onon, the range boundary goes westward somewhat to
the south of the southern end of [Lake] Baikal (it was recorded in
particular at the source of the Dzhida) and covers the Eastern Sayan
(sources of the Irkut and Oka).° Farther, the range enters Tuva (known
from Tanna-Ola) and, apparently, that more easterly part of the
Western Sayan which lies in the region of Us’ basin and northwest
of the city of Kizyl (Turan and Kurtushibinsk range—former Usinsk
region).

In Altai, the range occupies only the southern part of the coun-
try—Chuisk steppe and the region along the Chuya river, the area of
the upper Argut (Dzhassater-Yassater), and northwards to Uimon at
the source of the Katun [river]. To the west, the range enters the
upper Bukhtarma (Belaya Berel’) and the region of Lake Markakol’.
Farther, the boundary, enclosing the Tarbagatai and Saur, exits into
China.

The boundary again enters our territory as a small extension,
which irtcludes the Dzhungarsk Alatau and again exits into China,
entering our country once more along the Zailiisk Alatau. Further on,
it covers the whole Tien Shan including Kirgizsk (Aleksandrovsk)
range, Chatkal’sk and Talassk ranges and the ranges bordering the
Fergana valley on the north; the Alai system and Pamirs. Usually, its
occurrence is only documented for the East Pamir. This species prob-
ably exists also in the West Pamirs; however, information concerning
the Pamirs, as in general about all the mountains of Middle Asia, is
extremely rare. Apparently, the animal is absent in the Turkestansk
and Zeravshan ranges, as well as in the Gissar. From Pamir, the
boundary passes into Afghanistan (Kafiristan) and Kashmir.

The defined line of the northern limit of the range of the red wolf
is ill-determined. In many places, the animals are either extremely

In the eighteenth century, it was recorded considerably further north—in the
upper Lena (Pallas, 1811—“‘e superiore regione Lenae fl. albatus, vidi”). These state-
ments of this almost impeccable author are hardly simple errors (it is characteristic
that Pallas constructs the phrase in the plural). Probably, at that time, the animals were
distributed, perhaps transient, considerably more to the north than in our time or even
in the nineteenth century. In early times, in such places mountain sheep (Ovis
canadensis*—see vol. I), apparently existed.

*Now usually referred to O. nivicola—Sci. Ed.

rare or they appear fortuitously at intervals of several, ог even many
years. With respect to some areas, there are only old reports, uncon-
firmed in recent times. Lastly, information about some territories,
especially those such as Trans-Baikaliya are often contradictory. The
picture described here, discounting chance circumstances, is explained
not only by natural instability at the [range] limits but also popula-
tions at the edge of the range. Long-distance movements and
displacements, at least in some parts of the range are a characteristic
tendency for the red wolf (see below).

The red wolf is more common, and more or less regularly en-
countered, in the Ussuri territory, mainly in its southern part; rarer,
but also appearing more often than in other places (except southern
Ussuri territory), in the Tien Shan. Judging by data from old authors,
populations of red wolf within our borders considerably decreased
during the last 100—200 years. Apparently, the range has also con-
tracted, in any event the area more or less permanently inhabited.’

Geographic Range outside the Soviet Union

This extends from the northern limit as defined, southward through
the whole of Central and South Asia including Indochina and Ma-
lacca, Sumatra and Java and the whole of Hindustan, except its
northwestern desert sections and the extreme south. Here, the range
extends to Kurg (about 15° N. Lat.) and the Nilgiri Hills (11° N. Lat.).
Absent on the Japanese islands, Taiwan, Hainan, Borneo and Ceylon.
Information concerning its occurrence on Borneo, repeated in recent
literature, is undoubtedly mistaken. The red wolf is, apparently, ab-
sent in Iran also. Several suggestions (Byalynitskii-Birulya, 1912;
Ognev, 1931), based on data about Iranian Baluchistan (Zarudnii,
1931), were not justified.

Within the outlined range, the red wolf has in places been exter-
minated or crowded out by humans (East China and some parts of
India), and in other places it disappeared over significant expanses
without human interference. Such places include, apparently, some
parts of Tibet (in the Himalayas, around Lhasa in northeastern Tibet,

’Range according to data of Pallas, 1811; Radde, 1862; N. Severtsov, 1873;
Kashchenko, 1900; Bikhner [Biichner], 1902; Polyakov, 1914; Zarudnyi, 1915;
Tugarinov, 1916; Gassovskii, 1927; Kashkarov, 1927; Ognev, 1931, 1940; Yanushevich,
1952; Kuznetsov, 1948, 1948a; Sludskii, 1953, 1939; Shnitnikov, 1936; Spangenberg,
1936 and other authors.

392

578

in the Nan Shan; it is present in northern Kashmir—Ladak, Gilgit and
Chitral), desert and steppe areas of Inner Mongolia (present in mon-
tane sections of Dzhungaria and Kashgaria), some (steppe) parts of

northeastern China, eastern and central parts of the Mongolian Re-

public etc. Generally, the species is characterized by its peculiar
movement—irregular migrations—and also translocations within the
range. In some places in the center, and not only on the northern
edge, of the range, individual populations may move into other ter-
ritories where they had long been absent and then abandoned their
territory after several years. This is possibly connected with the ex-
termination of the large game by the wolves themselves (V.H.).

Geographic Variation

Geographic variation of the red wolf is, apparently, well manifested.
Usually, 11 subspecies are recognized (Pocock, 1936; 1941; Ellerman
and Morrison-Scott, 1951), which is, undoubtedly, exaggerated. In
our country, 2 subspecies may be accepted (Afanas’ev and Zolotarev,
1935). They are real, but their characteristics, due to collection
materials scarcity, cannot be considered well established.

1. Ussuri red wolf, С. a. alpinus Pallas, 1811.

General color tone of winter fur is intense rusty-red. Head dor-
sally, and outer side of ear, brownish-rusty; whole dorsal surface of
neck is same color, with blackish-brown highlights, shoulder and
entire upper surface of back brownish-rusty with black-brown high-
lights, outer side of legs rusty-brown color, lower side of body and
inner side of legs with noticeable yellowish tint.

Facial part of skull relatively narrow, forehead convex, nasal
bones elongated.

Dimensions comparatively large—greatest length of skull 180—
200, averaging 189 mm; length of nasal bones averages 66 mm.

Eastern part of the range in our country—from Pacific Ocean to
East Sayan. Western limits are not accurately known, but the Altai is
apparently not included in the range.

Outside the USSR, in the Mongolian Republic, northeast China,
and from eastern China southward, apparently, to Gansu, Sichuan and
Kam. Southern and western limits of distribution are not precisely
known.

Well differentiated, apparently the largest form, intense colora-
tion.

579

2. Tien Shan red wolf. C. a. hesperius Afanasjev et Zolotarev,
1935 (synonym jason).

General color tone of winter fur lighter, with weakly developed
rusty-red tints. Head dorsally, and outer side of ears reddish-straw
color, dorsal surface of neck dirty-white; along dorsal surface from
ears to shoulders runs narrow band of sandy-yellow color, shoulders
and entire upper surface of back faded yellow-brown color, outer
surface of limbs light sandy-yellow, ventral side of body and inner
side of limbs have no, or weakly developed, yellowish tint.

Facial part of skull relatively wide, forehead weakly convex, and
nasal bones shortened.

Dimensions relatively small—greatest length of skull 171.0 to
194.0, averaging about 180 mm; length of nasal bones averages about
57 mm.

Western part of range in USSR—Alltai, Tien Shan and, appar-
ently, Pamir.

Outside the USSR, the range of this form is unclear. Apparently,
it is distributed in China—along the Tien Shan and along the system
of ranges of the Kun Lun to Kukunor (Afanas’ev and Zolotarev,
1935). It is possible that its range is wider, and occupies the south-
western part of the [species] range (Kashmir, see below).

A form, well-differentiated from the nominal, but, apparently,
very close, if not identical with, that occupying Kashmir and West
Tibet.

Beyond the borders of the USSR, the following forms are usually
recognized (Pocock, 1936; Ellerman and Morrison-Scott, 1951, and
others):

1) С. a. dukhunensis Sykes, 1931—India south of the Ganges;
2) С. a. primaevus Hodgson, 1833—-Kumaon, Nepal, Sikkim, Bhutan;
3) C. a. lepturus Heude, 1892—China south of Yangtze-Kiang; 4) C.
a. infuscus Pocock, 1936—Tenasserim; 5) C. a. fumosus Pocock,
1936—western Sichuan; 6) C. a. laniger Pocock, 1936—Kashmir and
southern Tibet to Lhassa; 7) С. a. adustus Pocock, 1941—Upper
Burma, Indochina; 8) C. a. sumatrensis Hardwicke—Sumatra; 9) C.
a. javanicus Desm., 1920—Java.

It is quite evident that some of these forms are unfounded, for
example, those described from the southeastern part of the species
range. Probably, within the species limits, are not more than 4—5, and
not 11, forms.

393

580

In geographic variation of the species, Bergman’s rule is, appar-
ently, manifested quite well—compared to large animals from the
northeastern part of the range (nominal form), animals from the south-
eastern are noticeably smaller (V.H.).

Biology

Population. Extremely rare in the West Pamir, in Alai and western
Tien Shan (in Fergansk and Chatkal’sk ranges, Talassk Alatau,
Susamyrtau, Kirgizsk range), the northern (Zailiisk Alatau, Kungei
Alatau) and central Tien Shan (Atbash’ range, Tersk Alatau and
others), where it is caught individually, and not each year. In the 30’s
of the present century, in Kirgizia, | to 3 skins were tanned and also
not each year (Shnitnikov, 1936). From 1922 to 1956, a total of only
6 skins were tanned there (D. Dement’ev et al., 1956). Somewhat
more often, it is met with in the Dzhungarsk Alatau and its eastern
spurs, the Tarbagatai and Saur. In Tarbagatai, at the beginning of the
20th century, it was common (Plotnikov, 1912). In Saur and in the
eastern spurs of the Dzhungarsk Alatau, for instance, in the Alakul’sk
region, up to 6 red wolves were captured in a season in various years
(Khakhlov, 1928; A.A. Sludski1).

It is extremely rare in southern Altai (Kashchenko, 1899; Polyakov,
1914; Berger, 1946; Nasimovich, 1949). In the middle of the last
century in the Altai, it seems it was encountered somewhat more
often. In the ranges of the West and East Tannu-Ola and East Sayan,
it is rare (Radde, 1862; Solov’ev, 1921; Shukhov, 1925; Podarevskii,
1936; Yanushevich, 1952). In the Sayan and Okinsk territory and
Tafalaria, where in past years it was still sometimes found in packs
of 5—10 individuals, only a few are obtained individually and not
every year.

The red wolf is rare in the mountains of the southern part of Pri-
Baikal (Dzhidinsk and other ranges), in Trans-Baikaliya in the ranges
lying in the upper Onon river, on the northern slopes of the Great
Khingan and in the southern slope of the Stanovoi range, as well as
in the Dzhagda and Bureinsk ranges, although earlier it was common
in this region, and in some places, abundant (Maak, 1861; Radde,
1862). It is more often encountered in the Little Khingan and in its
foothills, and in the southern half of Sikhote-Alin. Earlier, it was
relatively common in the southern Ussuri territory, on the Yankovsk
Peninsula, and on the Iman (Ognev, 1931).

394

581

Fig. 148. Red wolves in winter. Specimens from China in the Moscow Zoo.
Photograph by A.P. Zhandarmov.

Habitat. In Middle Asia and Kazakhstan, the red wolf is met with
in summer in the alpine and subalpine belts at height of 2,500—4,000
m above sea level. Here, it occupies rocky heads of ravines, where
there are many Siberian ibex, or on the high-lying plateau—syrt, оп
which arkhar sheep are common. With the formation of deep snow
cover in the mountains, the red wolf follows after the ungulates,
moving to sunny southern slopes and to other light-snow regions. At
that time, it descends to the zone of spruce or juniper. In the lower
zones of the mountains, the wolves remain until the beginning of
summer, where at that time the females and lambs of the ibex, arkhar
sheep and roe deer dwell. In the spurs of the Dzhungarsk Alatau and
in the Tarbagatai, the red wolf was observed on gradual slopes inter-
rupted by rocky gorges, at a height of only 800—1500 m above sea
level. At such height here, ibexes, arkhar sheep and roes live.

582

In Altai, Sayan, Tannu-Ola and in Pri-Baikal, this animal is found
in summer in balds and high montane valleys with steppe vegetation,
ascending to a height of 1,500 to 2,000 m (Solov’ev, 1921;
Yanushkevich, 1952; А.А. Sludskii), but with the formation of the
deep snow cover, it departs to light-snow sections. In the mountains
of eastern Siberia, it was observed both in balds, and in taiga having
rock outcrops, along the lengths of rivers and badlands. It is attracted
to such places by the abundance of ungulates and the shallow, com-
pact snow cover. In the Far East, the red wolf lives not only in the
low mountains and foothills covered with Okhotsk taiga, or forests of
the Manchurian type with mixed deciduous species, but sometimes
descends to the seacoast.

Therefore, within the boundaries of the Soviet Union, the red
wolf is, in the greatest part of its range, a typical montane animal,
being that in the western half it lives, for the major part of the year,
high above sea level in alpine meadows and high-montane steppes. In
the eastern part of the range, this same carnivore is mainly an inhab-
itant of montane taiga, although here it ascends to balds; occasionally
it even appears along the seacoast. Everywhere it is restricted to
rocky places, in which it can take cover in case of danger.

The red wolf has a fluffy, dense fur with well-developed under-
fur, and its tail is also exceptionally fluffy. The ear tips are rounded
and are covered internally with dense wool. All these characteristics
testify to the adaptations of this animal as an inhabitant of severe
conditions.

Sometimes, red wolves appear in lands not familiar with them.
Thus, in 1901, on the Irtysh river, 70 km south of the city of Pavlodar
and about 150 km from the nearest mountains (Bayan-Aul), 2 red
wolves were caught on the grassy steppe. Earlier, these carnivores
were also observed there (Plotnikov, 1901, 1912). In the winter of
1954/55, a pair of red wolves appeared in the region of Smidovich
(west of Khabarovsk) in typical forest-steppe. Apparently, the ani-
mals moved to the forest-steppe from the Bureinsk range, avoiding
the deep snow cover (Sysoev, 1955, 1960).

In the Mongolian Republic, it lives in balds and foothill steppes.
On transiting from one group of mountains to another, it crosses great
expanses of dry steppes and deserts. It was observed on the strongly
desertified Tsagan-Bogdo range, which is rich in cliffs (Bannikov,
1954). In India, Burma, on the Indochinese peninsula, in Indonesia
and China, the red wolf inhabits the mountains from the alpine zone

395

583

(Himalaya and Tibet) to the foothills; preferring forest regions. In the
south of the range, it is common also on the plains and frequently
encountered on the seashore.

Food. Information about the food of the red wolf within the
Soviet Union is almost lacking. In the mountains of the Tien Shan,
and in the Tarbagatai and Saur, red wolves hunt Siberian ibex, arkhar
sheep, roe deer, maral [wapiti] and wild swine. The most frequent
prey of this predator is the ibex. It very rarely attacks domestic sheep.
In the Altai and Sayan, the list of animals that the red wolf hunts is
expanded to musk deer and reindeer. In eastern Siberia, the red wolf
preys, apparently, on roe deer, Manchurian wapiti, wild swine, musk
deer, reindeer, and in Primor’e, on spotted [sika] deer and goral as
well. Wolves appearing on the Yankovsk peninsula systematically
attacked spotted deer located in deer farms, and roe deer (Ognev,
1931).

In summer in the central Tien Shan, wolves made use, in their
diet, of plant in large amounts, mainly mountain rhubarb [Rheum].
This plant was always found in dens with pups, as if the wolves fed
it to the young, regurgitating the half-digested closed blossoms of the
rhubarb. The hunters, capturing live red wolves, fed these blossom to
the young captive animals (Shnitnikov, 1936).

In the Mongolian Republic, red wolves mainly hunt argali sheep,
more rarely, Siberian ibex (Bannikov, 1954). In China and India, red
wolves attack deer, sambars and muntjacs; antelope, mountain sheep,
wild swine, and even large wild oxen—banteng and gaur, as well as
domestic buffalo, horse and pigs. Packs of red wolves possibly dare
to attack bears, leopards and even tigers, sometimes driving them
away from their prey (Jerdon, 1874; Tate, 1947; Shaw, 1958 and
others).

Home range. Not established.

Burrows and shelters. No accurate data exist for the USSR. It is
known that for temporary lairs and whelping, they occupy small
natural caves, abandoned burrows of other animals, but rarely dig
their own. It has been suggested that sometimes several females whelp
in one den (India; Jerdon, 1874).

Daily activity and behavior. Red wolves hunt at any time of day
or night, but more often during the daytime. In the Tien Shan moun-
tains, they were repeatedly observed pursing prey during the day. The
red wolf runs more slowly than the jackal and red fox, but has great

endurance and can chase its victim for many hours. It is cautious and,

396

584

therefore, not easy to observe or hunt, but in attacking its prey,
especially when very hungry, it is daring and often kills its victim in
front of humans.

Except during the breeding period, the red wolf lives and hunts
in packs. In a pack occur 5 to 30, and sometimes more animals, but
usually 5—10. Thus, a pack of 5 animals was observed in late autumn,
1937, in the Kirgizsk range (Shnitnikov, 1936; О. Dement’ev et al.,
1956). In the Sayan, packs of 5—12 individuals were observed
(Solov’ev 1921; A.A. Sludskii). In spring, in Primor’e, a pack of 6
was noted and in winter up to 20 wolves (Ognev, 1931). This animal
hunts in China and India in packs of 5—10 individuals, rarely more
(up to 30) (Jerdon, 1874; Baikov, 1915; Tate, 1947; Shaw, 1958;
A.A. Sludskii; according to observations outside the borders of our
country).

Upon marking prey, the carnivore follows it for a long time,
driving it until able to get close to it. On running down a large prey,
they seize it by the thigh or side in the region of the groin, inflicting
wounds until the pursued animal falls. After bringing down a large
animal, the wolves sometimes bite its throat. Sometimes several wolves
run after the prey, while others run across to intercept. Cases are
known when red wolves drove the pursued animal to a precipice
where they killed it (Radde, 1862; Jerdon, 1874; Bourlieré, 1955).

According to G. Radde (1862), wolves upon sighting prey, utter
special whistling and hissing sounds, but this report needs new con-
firmation. Based on observations made in India, when pursuing prey,
they give a peculiar yelp from time to time, or else chase it in silence.

The idea that for a human to encounter a pack of red wolves may
be very dangerous (Radde, 1862; Sowerby, 1923) is a clear exaggera-
tion. In the Tien Shan, cases of attacks on humans are unknown. In
India, where this predator is common, attacks by it on humans are
also unrecorded (Ryabchikov, 1950); rarely, it attacks domestic ani-
mals there.

Seasonal migrations and transgressions. The sudden appearance
of a pack of red wolves at first one, and then another, place, and then
their disappearance for a long time, provides the foundation for sug-
gesting that, apart from the breeding period, they roam widely in
search of prey. With the appearance of their packs in a region rich
in ungulates, the carnivores systematically hunt, quickly dispersing
them, after which they are obliged to search for new hunting areas.
In the Tien Shan, red wolves do not live permanently in one place,
and they sometimes appear in those regions where they were previ-

585

ously not known, live there for some time and then disappear, but
after an indeterminate period of time they reappear. In the Chatkal
range, wolves always appear from the east, 1.е. from the main mon-
tane node of the Tien Shan (Shnitnikov, 1936). In Kazakhstan, cases
are known when red wolves appeared in the Chingiztau range (Karelin,
1841), whence they would pass from Tarbagatai, crossing in a straight
line for about 180 km. In order to reach Yamyshevskii village on the
Irtysh river (see above), they must have traveled in a straight line
about 600 km from the Tarbagatai. The wolves which appeared in the
former Smidovichesk region must have covered about 180 km, trav-
eling from the Bureinsk range.

Up to now in the Soviet Union, dens of the red wolf with young
were found only in the central Tien Shan. It is possible that they
generally do not reproduce at all in many regions within our country,
and that, of the red wolves which appear in the mountains of our
country, the majority only temporarily migrate to us. In the Mongo-
lian Republic, the appearance and disappearance of red wolves in
isolated free-standing ranges testify to their wide wanderings in this
country, during which they cross hundreds of kilometers through dry
steppes and deserts.

Reproduction. In the USSR, reproduction of the red wolf is
unstudied. According to observations in the Peking [Being] Zoo, its
rut occurs in January—February, and whelping in April. The period of
pregnancy is 60 days. In the course of a year it bears one litter. In
a litter of wolves in Peking Zoo, there were usually about 5 pups and
up to 9 (Krumbigel, 1954; Shaw, 1958). Apparently, larger litters
may occur, since the female has from 12 to 14 teats. The newborn
pup has a dark brown color.

Growth, development and molt. Not studied.

Enemies, diseases, parasites, mortality, competitors, and popula-
tion dynamics. Enemies include gray wolf, tiger, leopard and snow
leopard. These carnivores, at the same time, are manifested as com-
petitors. Diseases and parasites are unstudied. Population dynamics
are not clear. It is only known that, sometimes, red wolves become
more common in one or another place, a fact which is, however,
associated with their migrations from other regions (see above).

Field characteristics. Dimensions as in the juvenile gray wolf in
October, but with shorter limbs. The tail is longer with a fluffy end
which reaches the ground; in winter, covered with long wool, similar
to the fox. The animal does not lift the tail above a horizontal line.
Thanks to the ocherous-reddish-ginger color of the fur, the carnivore

586

397

is touted as “beautiful”’* from a distance, while the gray wolf, some
distance from the observer, looks almost white. The voice resembles
the yelp of a frightened dog and is not similar to a bark. The red wolf,
in contrast to the gray, does not howl (D. Dement’ev et al., 1956)
(A.S.).

Practical Significance

A fur-bearer, but because of its small number, it has no value in fur
tanning of the Soviet Union. It is not present in the fur standards and
its skin is accepted as that of gray wolf under the name “half wolf”.
Sometimes, skins of this animal are accepted as dog.

The winter skin of the red wolf with its long dense fur was highly
valued in the past by the Chinese, and the end of the 60’s of the
previous century in the Ussuri territory, they cost 3—4 silver rubles
(Przheval’skii, 1870), while at the beginning of the current century in
Manchuria, its price reached 8 rubles (Baikov, 1915). At the present
time the price of a prepared skin of the red wolf is from 1 to 4 rubles.
In Semirech’e, dokha [fur-coats] from the skins of the red wolf are
considered the warmest and are very costly.

Because of its small numbers and its cautiousness, it does not
harm livestock, although red wolves infrequently attack spotted deer
kept in parks (Menard, 1930; Ognev, 1931).

P.S. Pallas (1811) who first described the red wolf, put forth the
suggestion that this carnivore participated in the formation of some
domestic dog breeds; this idea is lacking in any foundation.

Taking into consideration that the red wolf is extremely rare, and
thanks to its small numbers causes no harm to the hunting economy
and livestock breeding, its special pursuit is not desirable. Until now,
hunting has been permitted throughout the year, and in Kirgiziya, a
bounty of 50 rubles is paid for each animal taken (А.З.).

Family of Bears
Family URSIDAE Gray, 1825
Carnivores of a generalized type, of large and very large size (this

family comprises the largest species of the order); they are heavy and
clumsy in built.

* The Russian phrase “Krasnye zver’” means “fine animal’—Sci. Ed.

398

587

Plantigrade, with broad, massive and short hands and feet. While
moving (at least our bears), they usually rest not on the whole front
limb but only on the anterior part of the fore limb only (digits and
anterior part of the metacarpal region). When moving faster, the
animals also do not rest on the entire hind limb (this is easily seen
in their tracks). In this way, they are, to a certain extent, digitigrade
animals. The bear rest upon the whole surface of the foot only when
moving very slowly or while standing.

Five digits, on anterior and posterior limbs of almost similar
length; all digits touch the ground. Their pads form only a slightly
curved line—pads of fingers II, Ш and IV lie only very slightly
anterior to the line on which are found pads I and V. Claws not
retractile, very long, curved in a sickle-shape and compressed later-
ally; on anterior limbs they are significantly, sometimes twice, as
long as on posterior limbs.

On lower surface of fore- and hind-foot, on anterior part of feet
themselves, except for digital pads there are transverse naked areas
(plantar and palmar callosities), and also bare areas behind them. In
different species, their relative size and form are very different; on
hind foot, they are usually larger. In some cases, entire lower surface
of fore- and hind-foot entirely covered by a large callosity. Last
phalanges of digits not capable of much upward flexing and have
normal articular surfaces.

‚ Skull relatively large and massive, in majority elongated; facial
part elongated and strong, with relatively long jaws, or else jaw and
facial part are much shortened (Malayan [sun] bear, Helarctos).
Zygomatic arches strong, fairly broad and general skull outlines pro-
portional, or facial part very short and zygomatic arches very widely
divergent laterally (Malayan [sun] bear). Contours of skull sharp,
sagittal (main posterior part) and occipital crests usually well devel-
oped; at point of their divergence, a posterior projection is formed.
Line of upper profile more or less convex, its highest point usually
located somewhat behind supraorbital processes.

Profile of facial part of skull concave in posterior section. Orbit
relatively small, temporal fossa very large. Mastoid region wide, and
forms behind posterior base of zygomatic arch a well-defined later-
ally directed process in forms of a platform. Paroccipital process well
developed, located quite far from tympanic bulla. Osseous tympanic
bulla usually flattened, partition in internal cavity absent; auditory
meatus in form of elongated tube. Alisphenoid canal present. Bony

588

palate narrow and elongated, its posterior edge lying behind posterior
point of last molar. Posteriorly, posterior palatine foramen relatively
far posterior, and located at level of last molar. On lower jaw, below
angular process, is a small inwardly directed alveolar process.

Complete dental formula (typical) is СРМ = 42, in one

case, СРМ = 40, (middle incisors absent [sloth bear]) and

3
in another, equivalent to normal, very commonly a and Bo (reduc-
2

tion of second and third premolars in association with extreme
shortening of facial part), i.e., total of 38 and 34 (Helarctos). Even
in forms with complete dental systems (Ursus s.1.), as individual and
age-related characteristics, there is loss of some of the anterior three
premolars, especially the second upper and second and third lower.
Frequently, therefore, even in species with complete dental formulae

(brown bear), it may take the form: [5c PoM> = 34. Incisors form
a weakly convex line, lateral larger than [two] interior, sometimes
considerably; canines strong, broad at base. Entire cheek tooth row
and canines form almost straight line, hence both rows parallel, first
three premolars in both jaws small, conical, have only one root and
frequently absent (fallen out).

True carnassial teeth do not develop. Fourth upper premolar small
(smaller than first molar), usually having three blunt cusps—one
anterior and two posterior. It is remarkable that it has two roots and
is situated anterior to the infraorbital foramen and is the point of
maximum mechanical force. Upper molars large, with broad low
crowns without cutting elements on surface. First has four more or
less clearly defined cusps, separated by a middle longitudinal fissure.
Second upper molar larger than first, extending in longitudinal direc-
tion, with large “heel” and a broad longitudinal groove in middle; two
cusps are located on its outer side.

In lower jaw, fourth premolar small, conical in form; first molar
elongated, with three cusps on outer part and well developed heel,
usually carrying two cusps. Second molar more or less same length
as first, but broader. In middle, has longitudinal fissure, outlined by

399

589

marginal crests, in which are elements of two cusps. Third molar
smaller than others, with flat, slightly projecting cusplets, outlined by
low пт.

Praeputial orifice situated far anterior to scrotum. Penis long,
glans penis smooth, without spines. Os penis present. Size of bone
quite large, tapered on end, longitudinal groove absent. Anal glands
absent or extremely poorly developed. Tail very short, usually hidden
in fur and unnoticeable. Ears rounded, upright, in some species quite
large, in others hardly protruding from fur. Eyes small. End of nose
bare and lips, both upper and lower, large and movable.

Pelage long and shaggy, in northern and montane forms, thick;
except in one species (Helarctos), short and quite sparse. Color mono-
tone—white, black or brown of various tones—from golden or
yellow-brown to dark brown. Individual and geographic variation in
color very great in some forms (brown bear, Ursus arctos); in major-
ity of forms, color is stable. In one species (American black bear,
Ursus americanus), color is triphasic—black, brownish, and “Бше”.*

In some species, qualifying as a constant character is a white spot
on the chest (white-chested or Himalayan black bear, Ursus
tibetanus**, Malayan sun bear, Helarctos), or it appears in the nature
of individual deviation (brown bear; American black bear Ursus
americanus), sometimes developing as a light collar; in some forms,
a particular color of muzzle qualifies as a permanent character (light,
brownish—American black bear, Malayan bear), or a light ring around
the eyes (spectacled bear, Tremarctos), etc.

One pair of pectoral teats. In several species (brown bear), there
are three pairs, but only one pair of functional mammary glands. It
is possible that this also occurs in other species, but data on this are
lacking.

Sexual dimorphism in color and other characters is absent, but
males are larger than females. Age morphism, as a rule, is also
absent, or is weak (young brown bear although monotone, usually has
a light spot on chest or neck). Seasonal dimorphism is considerable
only in northern forms and is expressed in different length and thick-
ness of winter and summer fur. There is only one molt annually.

In general appearance, all species of the family are quite homo-
geneous, and represent insignificant deviations from one bear type.

*A whitish phase is also known—Sci. Ed.
**Sic; should be thibetanus—Sci. Ed.

400

590

All have a weighty general structure, which is emphasized by ab-
sence of tail, and in the majority, long shaggy fur; legs are short with
large, sometimes with extraordinarily [ong curved claws; head is large,
heavy, usually furry, with small, deeply set eyes, withers in the majority
are high. Depending on development of climbing ability, whether
they are more or less arboreal, or, on the contrary, purely terrestrial
in way of life, the degree of massiveness of anterior and posterior
parts of the body differs. In a purely terrestrial form (polar bear), the
posterior part of the body and muscles of the hind legs are especially
developed; and in a good climber (white-chested bear, sloth bear,
Malayan bear), anterior part of the body and muscles of anterior
extremities are especially developed.

Differences in size of different species are slight. The largest
species (polar bear, brown bear) in their largest forms have a maxi-
mum body length of 230 to 300 cm; a height of 135 cm at the
shoulder, and a weight of 500 to 700, and even, apparently, 1000 kg
(brown and polar bears). The smallest species (Malayan bear,
Helarctos) has a body length averaging 115 cm and a weight of 50
to 100 kg (Pocock, 1941). Therefore, the ratio between the weight of
extreme forms constitutes |: 10 or less.

Bears are mainly associated with forests of the plains and moun-
tains of temperate latitudes and tropics, and often in forestless montane
regions; only one species inhabits the shores of the Arctic Ocean and
on arctic ice, being a semiaquatic animal. In a vertical direction, its
distribution is from sea level to alpine meadows and the highest
plateaus of Central Asia.

The bear usually moves slowly with heavy steps; in case of
danger some run very rapidly; some species swim well. All except
the polar bear climb trees well (brown bear only when young); some
represent quite highly developed specialized arboreal forms (sloth
bear, Melursus; Malayan bear, Helarctos), with some particular
adaptations (structure of the fore-limbs and others). They are solitary
animals, remaining in families (females with young, sometimes the
male). In the case of food, they are omnivorous animals; some are
more specialized on plant foods (grass, berries, nuts), others on ani-
mal, feeding in particular on vertebrates or invertebrates including
insects (bees, ants, termites, etc.). One species (polar bear) feeds
exclusively on meat of large animals. The majority of species are
quite strictly sedentary, while others (polar bear), roam widely the
entire year (males) or part of the year (females with young).

591

In some species, all, or almost all populations (except extreme
southern) hibernate (brown bear, white-chested bear, American black
bear, Ursus americanus); in one species, only pregnant females live
in dens (polar bear). Some species do not hibernate. They are
monogamous*, however, they do not form permanent family pairs
and the male does not take part in the care of offspring. There are 1,
2, or rarely 3, young.

They are mainly nocturnal, partially diurnal, or active throughout
the day/night. Vision is weakly developed, hearing and smelling are
very sharp. Mentally, it is one of the highly developed groups of the
order; some forms in particular respond well to training.

The range of the family is vast and, in some respects, very unique.
It is divided into two parts; the chief of which includes both land and
open sea. The principal mass of the range occupies North America,
Eurasia, the Arctic basin and a small area in northern Africa. A
isolated section of habitat is found in South America. Bears are ab-
sent in almost all of Africa and in Australia, as well as on the greater
part of the island region between Asia and Australia.

The established southern border of the northern part of the range
in the New World covers the islands of Saint Lawrence, Saint Mat-
thew**, Hall**, Nunivak**, and Pribilof**, the Alaska Peninsula
(Bering Sea), and the eastern islands of the Aleutian ridge, westwards
to include Unimak; then goes along the western coast of North
America, including islands of Kodiak, Queen Charlotte, Vancouver
and others and the northern part of the California peninsula, and
crosses the mainland along Mexican territory. Here, the border goes
southwards along the Sierra Madre system to the region of Durango
and reaches the Tropic of Cancer, extending even a bit farther south.
The border extends farther, along the coast of the Caribbean Sea to
the Florida coast and northwards along the eastern coast of North
America. Covering Newfoundland, it directs itself to the water at the
southern tip of Greenland.

In the Old World, the limits of the range covered all of Europe,
from northern Iceland and Great Britain, though without the islands
of the Mediterranean Sea; the Atlas region in North Africa, Asia
Minor, Syria and Palestine, Iraq, Iran, Pakistan, India, Ceylon,

*Sic; most, if not all, bears are presently considered promiscuous, not forming
a pair bond of more than a few weeks—Sci. Ed.
**No records of bears from these islands—Sci. Ed.

402

592

Indochina, Malacca, Sumatra and Kalimantan (Borneo). Farther, it
extended along the eastern coast of Asia, including Hainan, Taiwan,
(absent on the Ryukyus), the Japanese islands, Sakhalin, Shantar is-
lands, southern (Kunashir and Uturup) and northern (Shumshu and
Paramushir) Kuril islands (absent on the remainder), Kamchatka and
Karagin Island. On the Commanders they are absent. The range
extended northwards to the high latitudes of the Arctic Ocean—
apparently to the pole.

Within the outlined regions, there is no occurrence of the family
over significant areas due to the unfavorable natural conditions (steppe,
deserts, open tundra); in several regions they appear only accidentally
(Iceland), or during migration, and do not reproduce (pelagic parts of
the Arctic Ocean). In large territories they were exterminated (part of
the United States, British Isles, some parts of Western, Central and
Eastern Europe, North Africa, Japan and others).

In South America, the range occupies the higher (to 3,000 m)!
parts of the Andes in western Venezuela, Colombia, Ecuador, Peru,
and western Bolivia. Here, it was also reduced.

Everywhere, reduction of the range of the family represents only
the result of their direct pursuit and destruction by humans. Based on
all the characteristics of this unique family, it is an entirely recent,
ecologically flexible and viable group.

Within the limits of the superfamily* and groups of Canoidea,
this family is well isolated and sharply differentiated from the others.
For a long time, the composition of the family included the genus
Ailuropoda (bamboo bear, or giant panda), producing a famous lack
of clarity in determining the limits of the family. After excluding this
genus from Ursidae and placing it in Procyonidae (raccoons), or
grouping it together with Ailurus (panda, cat-like bear) in a special
family, the limits of the described family became fully legible and
natural**.

As a whole, the species of this family represents carnivores of a
poorly specialized type. This is expressed both in characteristics of
the skull and the dental system, and also in such characters as plan-
tigrade limbs. In regard to its position among the recent families of

'As was recently clarified, references to northern Chile were mistaken (Cabrera,
1957).

*In Russian original, mistakenly called “order’—Sci. Ed.

**Scientific opinion has recently swung back to the earlier view—Sci. Ed.

‘(GamMdoH `9’Л) “AND snjpuso sojaipwasy ‘леэа pajoejoads ay} jo
эЗиег—воцэшу YINOS ш эий PI[OS “eISY ш ‘1 51540] ‘51894 эп JO $пиэ8 dy} JO 1эр104 WaYINOS pajonssuode1—oul] рэцзер ‘эЗие! JO
1эр104 изэц}по$ SMOYS SUT] PHOS “POM PIO OU) Ul рие вомэшу чном и! ‘эер!з1 ‘yt леэа oY} JO эЗие1 рэзэп45и0ээ ‘6 “SIA ТОР

594

the order, contemporary Ursidae reveal a significant relationship to
the recent Procyonidae (raccoons). This is also partially indicated by
the tale concerning the giant panda (Ailuropoda). Its relationship to
modern Canidae (wolf family) is, apparently, not too close.

As regards its origin, the family is somewhat less ancient than
Canidae, and the earliest forms appear in the middle or lower Mio-
cene (Ursavus, Europe). Usually, the origin of the family is connected
with the extinct subfamily Amphicynodontinae (Cynodontidae) of the
family Canidae, which existed from the upper Eocene to the lower
Pliocene. Among this group, the nature of originating material that
gave rise to the recent Ursidae is usually given as Cynodon, Dinocyon,
Cephalogale and several others. The divergence of bears from these
truly flesh-eating carnivores was apparently determined most of all
by their adaptation to eating plant foods. The dental system has
changed correspondingly.

The development of bears proceeded, apparently, in Eurasia,
whence the earliest forms of Ursidae are known, and also from which
they are accepted as having derived. However, the old opinion, that
in the New World this family appeared only in the Pleistocene, is
wrong—from here representatives of the family are known from the
lower Pliocene and, perhaps, even from the middle Miocene (Ursavus).
It was never, one thinks, a family especially diverse and rich in
species.

In internal systematic structure, this family is sufficiently homo-
geneous that subfamily groupings are not noted in it, and in this all
investigators agree’. In the handling of generic grouping, however,
there is no agreement, and several authors even in our day put each
species into an independent genus, or for 7 species, 6 genera (Pocock,
1941; Simpson, 1945; Ellerman and Morrison-Scott, 1951). This is an
obvious exaggeration and the actual number of genera is undoubtedly
less. This extreme point of view, and in general the strong division
into genera, is explained by the fact that bear species are sharply
differentiated from each other, and because morphological degrees of
difference are great and clear. Among recent bears, forms whose

Тре recent suggestion (Thenius, 1959; Thenius and Hofer, 1960) to divide the
family into 2 subfamilies—Tremarctinae (American spectacled bear—one genus and
species) and Ursinae (all other bears), based on paleontological considerations is
incompletely argued, and is not accepted (see further, characteristics of the genus
Ursus).

595

403 species independence might be doubtful and could be grouped to-
gether, are absent.’

In all, the family comprises 11—12 extinct, and 3—4 living gen-
era.* The recent fauna may include the following genera: Ursus
including Thalarctos, Euarctos, Selenarctos, i.e. brown, [Asian] black
and polar bears, as well as the American black bear; Melursus (sloth
bear); Helarctos (Malayan bear) and Tremarctos (spectacled bear).
The latter genus is closely related to the genus Ursus and, perhaps,
may deserve to be combined with it.

Melursus 13 also closely related to Ursus. Helarctos is considered
to be the most sharply deviant form—all remaining bears are closer
to each other than Helarctos is to any of them. It seems as if to be
contrasted to the others. Perhaps it might be more natural to divide
the family into two genera—Helarctos with one species (H.
malayanus), and Ursus, including all the remaining. At the same
time, one may note a certain type of relationship between the genus
Ursus s. |. and the genus Helarctos, through the white-chested bear.
(U. thibetanus) (see description of white-chested bear), which make
this proposed division not excessively sharp. Within the genus Ursus
it is acceptable to create subgenera (see below). Helarctos represents
the most specialized form (arboreal). The genus Ursus must be con-
sidered as a less specialized group; however, within it the degree of
specialization is, in its turn, quite diverse (see description of genus).°

In the family there are, in all, 7 species, in the genus Ursus—
4 (see characteristics of genus), in genus Tremarctos—1 (T. ornatus
Cuvier, 1935—spectacled bear of South America); in genus Melursus—
1 (М. ursinus Shaw, 1880—South Asiatic sloth bear) and in genus
Helarctos—1 (H. malayanus Raffles, 1822—South Asiatic Malayan
[sun] bear).

The practical significance of the species in the family is small. In
different places, some cause definite harm to animal breeding, which
is, however, inconsiderable. Almost all of them serve as objects of
sport hunting, several of which in this sense are of great value (brown
bear). Some have commercial importance for local needs or provide
marketable skins (polar, brown bears). Numbers of a series of species

The unlimited division of North American bears suggested by Merriam (1918),
and which is still partly adhered to, is set to one side. It lies outside scientific zoology.

“According to Simpson, 14—18 extinct and 6 living.

°On the contrary, Thenius and Hofer (1960) consider Helarctos a less specialized
form, and the brown bear, a specialized form.

404

596

have greatly decreased in connection with their pursuit, and hunting
of individual species in parts of their range is found to be prohibited
(polar bear). In some places, bears are considered as pests and are
liable to be destroyed.

In the fauna of our country, there is one genus—Ursus Linnaeus,
1758, which comprises 25% of the genera of the family, and 3 spe-
cies, 1.е., about 43% of the species of the family. Bears constitute
about 1% of the species of our fauna.

The range of the family covers almost the entire territory of the
Union except the steppes and deserts and parts of the tundra. They
are found on the Arctic Ocean.

They are game animals, objects of sport, as well as commerce.
One species (polar bear) is considered endangered; hunting of the
others is not restricted or is only restricted in a part of the distribution
area (V.H.).

Key for Species Identification of the Bear Family

1(2). Color of fur white or with gray-yellow tint, bases of hairs white.
Hairs on bottom of foot strongly elongated and much longer
than on upper side. Upper molar teeth (last two teeth in tooth
row; anterior one is first, and posterior is second) not large, total
length less than distance between the anterior (first molars) of
the right.and, left sidesy(Fig: 150) т
к. sets cea ao Polar bear, Ursus maritimus (p. 678).

2(1). Color different; bases of hairs black or brown. Hairs on bottom
of foot not elongated, and in length not different from hair on
upper side of foot. Upper molar teeth are large and elongated,
their total length more than distance between anterior (first)
molars of right and left sides.

3(4). Color from straw-brown to dark-brown. No sharply outlined
clear white area on chest. On lower surface of fore foot, two
large bare areas in addition to bare digital pads. Distance from
middle of line joining postorbital processes to anterior end of
premaxillae equal or nearly equal to distance from above-men-
tioned line to posterior point of occipital crest. Anterior edge of
last upper molar lies anterior to anterior lower edge of orbit. .
о oul Brown bear, Ursus arctos (p. 601).

4(3). Color is pure black or with very weak brownish frosting. On
chest, sharply outlined clear white, half-moon-shaped field or,

1758.

1825:

1825.

1855.

1864.

1865.

1896.

1898.

1901.

LOL 7:

1923.

Si,

in any event, extending in transverse direction. On lower sur-
face of fore foot, in addition to digital pads, one large naked
area, occupying whole surface of foot. On skull, distance from
line connecting postorbital processes to anterior end of premax-
illae considerably less than distance from above-mentioned line
to posterior point of occipital crest. Anterior edge of last upper
molar lies at level of anterior lower edge of orbit...... White-
chested [Asiatic black] bear, Ursus thibetanus (p. 713) (V.H.).

Genus of Bears
Genus Ursus Linnaeus, 1758

Ursus. Linnaeus. Syst. Nat. Ed. X, 1, p. 47. Ursus arctos
Linnaeus.

Thalarctos. Gray. Ann. of Philosophy, N.S., 10, p. 62, July
1825. Thalarctos polaris Gray = Ursus maritimus Phipps.
Thalassarctos. Gray. Ann. of Philosophy, N.S., 10, p. 339,
November, 1825. Substitute for Thalarctos Gray.
Tremarctos. Gervais. Hist. Nat. Mammif. 2, p. 20. Ursus ornatus
Cuvier. (Name used for Ursus t{h]ibetanus).

Myrmarctos. Gray. Proc. Zool. Soc. London, p. 694.
Myrmarctos eversmanni Gray = Ursus arctos Linnaeus.
Euarctos. Gray. Proc. Zool. Soc. London, 1864, p. 692. Ursus
americanus Pallas.

Thalassiarchus. Kobelt. Bericht. Senkenberg. Naturf. Ges.
Frankfurt am Main, p. 93, Substitute for Thalarctos Gray.
Ursarctos. Heude. Mem. Hist. Nat. Emp. Chin., 4, pt. I,
p. 17. Ursus yesoensis Lydekker = Ursus arctos Linnaeus.
Melanarctos. Heude. Mem. Hist. Nat. Emp. Chin., 4, pt. I, p.
18. Melanarctos cavifrons Heude = Ursus lasiotus Gray =
Ursus arctos Linnaeus.

Selenarctos. Heude, Mem. Hist. Nat. Emp. Chin., 5, p. 2. Ursus
thibetanus Cuvier.

Arcticonus. Pocock. Ann. Mag. Nat. Hist., 20, p. 129. Ursus
thibetanus Cuvier.

Mylarctos. Lonnberg. Proc. Zool. Soc. London. p. 91. Ursus
pruinosus Blyth. = Ursus arctos Linnaeus (V.H.).

405

598

Dimensions large and very large.

Form and size of naked areas (“callosities”) on lower surface of
fore- and hind-feet vary in different species—in extreme cases, they
occupy nearly their entire surface or, on the contrary, lower surface
of foot almost entirely covered with hair.

Skull large and massive, with strongly or moderately developed
facial portion and strong, relatively long, jaws. Zygomatic arches
moderately broad and general outlines of skull proportional.

Зи 42

Dental formula а = 42.

Middle upper incisors always present, but first three premolars
weakly developed and some of them frequently absent. They
are usually shed with age, but sometimes do not develop a
definitive system at all. More often, second and third premolars ab-
sent, sometimes also first. Fourth premolar always present. There-
fore, dental formula individual in nature; in part age variation may

. В tae) ЗА 2. sea)
take the form I—C—P—M—=38; 1-С-Р-М- = 34 and even
Е Sie VAD ATA

СРМ = 30, i.e. deviating far from normal limit of family.
Relative size of molars varies somewhat in different species.

Ears vary in size—of moderate length, distinctly protruding from
fur, large and broad; or relatively small, slightly protruding from fur.
Size of bare portion of nose and dimensions of lips vary in different
species.

Pelage dense and long. Color of two species is very uniform, in
one white with yellow highlights, the other pure black (polar and
white-chested Asian black bear); one species is very variable as re-
gards color—both individually and geographically it varies from dark
brown to straw yellow (brown bear); one species (American black
bear, U. americanus) has triphasic coloration, black, light brown and
“blue”*. In one species, as a permanent qualitative feature, there is
a large white spot on the chest (white-chested bear); lightening on the
chest, sometimes a light collar appears in the form of an age character
in young animals, and sometimes as an individual variation in brown
bear.

*A forth, white, phase also occurs—Sci. Ed.

406

599

In the majority of forms, the posterior part of the body is highly
developed and more massive; in one species—the anterior (white-
chested bear). The genus comprises the largest forms in the family
and order (see above, characteristics of the family and below, data on
individual species), and species of moderate size, with a long body,
about 180 cm and weight not more than 250 kg (American black
bear).

The species of the genus are distributed in forest regions, in part
in montane forests or in treeless mountains and plateaus, sometimes
even in deserts, and extending into the tundra; one species is linked
to ice and coasts of the Arctic Ocean, being a semiaquatic animal.
Some species climb trees well, but in a restricted sense, these arbo-
real forms represent several other genera. They are omnivores,
preferring plani forms, [but] one species (polar bear) is exclusively
carnivorous. They are sedentary forms, [but] one species roams widely
(polar bear). All species hibernate in winter.

The range of the genus (reconstructed) occupies the part of the
general range of the family, being located in the Arctic Ocean,
Greenland, North America, Europe, in northwestern Africa and in
Asia excépt its extreme southern parts. Here, the southern range
border of the genus crosses through Asia Minor, Palestine, Syria,
Iraq, southern Iran, montane regions on the right bank [west] of the
lower and middle Indus [river] along the Himalayas, and covering all
Indochina, except the southern and middle parts of Malayan Penin-
sula (for more details, see description of range of brown and
white-chested bear). During the past century, the range was strongly
reduced, and members of the genus are already absent in a large
expanse. This applies to the greater part of North America, to Western,
Central and Eastern Europe (European part of our country). Reduction
has occurred locally in other parts of the range, but to a lesser degree.

In the genus there are 4 species—U. arctos Linn., brown bear; U.
thibetanus Cuv., white-chested, or [Asian] black bear; U. americanus
Pall., American black bear or baribal; and И martimus Phipps, polar
bear.

Contemporary representations of the species content of the genus
given here were established only recently and are more or less widely
accepted. However, in literature up to now, echoes of mistaken incor-
rect inferences still occur. The specific unity of all brown bears of
Eurasia and also North America is sufficiently evident, and this was
directly stated quite long ago not only in Russian, but also in the

600

Western European literature. However, even in 1931, when the con-
temporary broad concept of the species had not yet attained currency
among our mammalogists, in the USSR the genus Ursus comprised
six species (arctos, piscator, mandschuricus, jessoensis, pamirensis,
pruinosus; Ognev, 1931). For Central Asia two species occurred
(arctos, pruinosus; G. Allen, 1938).

In particular, the systematics of the American bears were misrep-
resented, and even now are often mispresented. For only the American
part of the range, Merriam (1918) separated 77 species, several with
subspecies (total of 84 forms); one of these species was even sepa-
rated in a special genus (Vetularctos). The “concept” of Merriam
clearly lies outside of zoological science and generally cannot be
taken into consideration, or has purely nomenclatural, even negative,
significance. The species unity of American and Eurasiatic brown
bears is, as was told, itself quite evident and was, more than once,
confirmed in literature. Nevertheless, this “system” is still found re-
flected up till now in reports concerning the fauna of America
(Anderson, 1946; Miller and Kellog, 1956).

Within the limits of the genus, the terrestrial form, the brown
bear must, apparently, be considered the least specialized “general-
ized” one. The American black bear (U. americanus) is closest to it.
They may be taken as one subgenus, Ursus. Another direction of
specialization—greater climbing ability—is represented by the white-
chested, or Asian black bear, which may be separated into a distinct
subgenus, Selenarctos. The polar bear represents another direction of
specialization related to water. They are usually accommodated under
the subgenus (genus) Thalarctos. At the same time, if we leave to one
side color and several other external features, the similarity between
polar bear and brown is found to be very great, and a series of authors
deny the basis for separation of polar bear into a special subgenus.
The proximity of these species is also indicated by occasional cases
of hybridization.

Different authors, more than once, confirmed the closeness of the
Asian and American black bears—U. americanus and U. thibetanus.
Some authors directly ascribe them to one genus (subgenus), Euarctos,
placing them opposite the brown bears—Ursus $. str. This point of
view was fully developed by G. Allen (1938). At the same time, as
is in part clear from material to be brought out later, U. thibetanus
itself represents a form quite strongly deviating from the purely “ter-
restrial” type of bear, while U. americanus, according to general

407

601

adaptive type—is a true “terrestrial” bear and is quite close to U.
arctos in skull structure. Not only is its combination with U. thibetanus
in one subgenus wrong, but also its separation in the subgeneric sense
from the brown bear—U. arctos.

In relation to the Asiatic black bear, the generic name 7remarctos
(Pocock, 1914) is also employed, which accentuates its proximity, in
several respects, to the South American spectacled bear (7. ornatus).
This, however, can hardly be accepted.

Within the boundaries of the USSR, there are three species: 1)
brown bear. U. (Ursus) arctos Linnaeus, 1758; 2) polar bear, U.
(Thalarctos) maritimus Phipps, 1774; and 3) the white-chested or
[Asian] black bear, U. (Selenarctos) thibetanus Cuvier, 1823. They
constitute about 43% of the species in the genus and about 1% of the
number of species in our fauna.

In the USSR, the species of the genus occupy a large part of the
country, except the Asiatic steppes, semideserts and deserts and part
of the tundra (reconstructed range). Ranges of all species have de-
creased.

They are game and economically exploited animals in part con-
sidered as pests and pursued under law. One species (polar bear) is
preserved by law (V.H.).

Subgenus of Brown Bears
Subgenus Ursus Linnaeus, 1758
BROWN BEAR
Ursus (Ursus) arctos Linnaeus, 1758

1758°. Ursus arctos. Linnaeus. Syst. Nat. Ed. X, 1, p. 47. Sweden.

1788. Ursus arctos niger. Gmelin, Syst. Nat. Ed. XIII, 1, p. 100,
Northern Europe.

1792. Ursus arctos griseus Kerr. Anim. Kingd., p. 188. Germany.

1820. Ursus arctos minor Nilsson. Skand. Fauna, p. 123. Very north-
ern part of Scandinavia.

°Synonomy of European and North European forms of bear representing a pure
literary and historical interest, given in short form. For details see Miller, 1912;
Ellerman and Morrison-Scott, 1951; Couturier, 1954.

408

602

1824.

1826.

1827.

1828.

1840.

1851.

1854.

1855.

1864.

Ursus collaris Cuvier et Geoffroy. Nat. Hist. Mamm., pt. 42,
pl. 212. Siberia.’

Ursus isabellinus Horsfield. Trans. Linn. Soc. Zool., 15, p.
334. Mountains of Nepal.

Ursus arctos brunneus Billberg. Synop. faunae Scand., p. 15.
Northern Scandinavia.

Ursus arctos myrmecophagus Billberg. Ibidem, p. 16. North-
em Scandinavia.

Ursus arctos formicarius Billberg, Synop. faunae Scand., ed.
2, p. 16. Renaming of U. a. myrmecophagus.

Ursus syriacus Hemprich et Ehrenberg. Symbol. Phys., 1, pl.
1. Lebanon.

Ursus cadaverinus Eversmann. Bull. Soc. Imp. Nat. de Moscou,
p. 11. Renaming of U. arctos.

Ursus longirostris Eversmann. Bull. Soc. Imp. Nat. de Moscou,
p. 11. Renaming of U. formicarius.

Ursus arctos var. normalis Middendorf. Verhandl. d. Minerolog,
Ges. zu St. Petersburg. Jahrg. 1850—1851, p. 74. Northeastern
Europe.

Ursus arctos var. meridionalis Middendorf. Ibidem, p. 74.
Caucasus.

Ursus arctos var. beringiana Middendorf. Ibidem, p. 74.
Northwestern America.®

Ursus arctos var. kamtschatica Middendorf. Ibidem, p. 74.
Nomen nudum. Obviously, Kamchatka.’

Ursus pruinosus Blyth. J. As. Soc. Bengal, 22, p. 589. Lhassa,
Tibet.

Ursus arctos aureus Fitzinger. Wiss.-pop. Naturg. 4. Saugeth.,
1, p. 371. Eastern Russia.

Ursus piscator Pucheran. Rev. Mag. Zool., 7, p. 192.
Petropavlovsk, Kamchatca.

Ursus arctos var. | normalis. Gray. Proc. Zool. Soc. London,
p. 682. Renaming of arctos.

7See “Note” in description of Eastern Siberian bear in the section “Geographic
Variation”. ;

Рог type locality of this form and nomenclatural significance of name, see
“Note” in description of Kamchatka bear in the section “Geographic Variation’.

*Pagination in- this work given according to separate reprint.

1864.
1864.

1864.

1864.
1867.

1873.

1833.

1883.
1897.
1898.
1898.
1913.

1916.

1924.

603

Ursus arctos sub-var. а. scandinavicus Gray. Ibidem, р. 682.
Scandinavia. |

Ursus arctos sub-var. с. rossicus Gray. Ibidem, р. 682. Nom.
nud. Evidently, Russia.

Ursus arctos var. sibiricus Gray. Ibidem, p. 682. Siberia.
Ursus arctos var. 2 grandis Gray. Ibidem, p. 684. Northern
Europe.

Ursus arctos var. 4 stenorostris Gray. Ibidem, p. 685. Poland.
Myrmarctos eversmanni Gray. дет, р. 695. Norway.
Ursus lasiotus Gray. Ann. Mag. Nat. Hist., 20, p. 301. “Inner
part of Northern China”. Located more exactly: eastern part of
Manchuria.

Ursus leuconyx Severtsov. Vertik., gorizont. raspr. Turkest.
zhivot. Izv. Obshch. Lyub. est. antrop. etnogr., 8, No. 2, p. 79.
Valley of upper Naryn, Tien Shan.

Ursus lagomyarius Przevalski. Przheval’skii Tret’e putesh. v
Tsentr. Asii [Third journey in Central Asia]. p. 216. Mountains
of Kuku-Shili, Eastern Tibet (35° N. Lat., 96° E. Long.).
Ursus hypernefes Przevalski. Przheval’skii, Ibidem, p. 216.
Substitute for U. lagomyarius Przev.

Ursus arctos yesoensis Lydekker. Proc. Zool. Soc., London,
p. 422. Hokkaido Island, Japan.

Ursus mandchuricus. Heude. Mém. Hist. Nat. Emp. Chin., 4,
р. 23—24, pl. 7. Near Vladivostok.

Melanarctos cavifrons. Heude. Ibidem, 5, р. 1. Tsitsikar, north-
eastern Manchuria.

Ursus arctos lasistanicus Satunin. Tr. Obshch. izuch.
Chernomorsk, poberezh’ya, 2, p. 27. Sukhumskii area.
Ursus arctos var. caucasicus Smirnov., Zap. Kavk. Muzeya,
ser. A, No. 4, p. 117. Passanaur on the Georgian Military
Highway (southern slope of Main [Caucasus] Range).

Ursus arctos arctos L. natio dinniki Smirnov. Ibidem, р. 122.
“Chatakh, Borchalinsk. u. [county], Tiflissk. guber. [gover-
nance]”.

Ursus arctos yeniseensis Ognev. Priroda i okhota na Ukraine
[Nature and Hunting in the Ukraine], 1-2, р. 112. Maina,
tributary of Ungut R[iver], vic. of Krasnoyarsk.

Ursus arctos baicalensis Ognev. Ibidem, р. 112. Kudaldy,
Trans-Baikaliya.

409

604

1924. Ursus arctos kolymensis Ognev. Ibidem, p. 112. Zabortsevo on
the Kolyma [River].

1924. Ursus yessoensis Ognev. Ibidem, p. 112. In place of yesoensis
Lyd. 1897.

1924. Ursus pamirensis Ognev. Ibidem, р. 111. РатизКи post, Pamirs.

1925. Ursus arctos persicus Lonnberg. Fauna och Flora, 1, р. 28.
Mezanderan, N. Iran.

1925. Ursus arctos smirnovi Lonnberg. Ibidem, р. 28. Northern slope
of Main Caucasus Range (V.H.).

Diagnosis

Dimensions large to very large. Color normally brown, of different
intensities, but individually, and in part, geographically very strongly
variable—from straw-yellow to almost black. On lower surface of
forefoot, in addition to five digital callosities (pads), there is a large
transversal anterior callosity and a separate small rounded posterior
one located nearer outer edge of foot. Facial part of skull relatively
long, last upper molar broad and elongated, considerably larger (nearly
double) than anterior; last lower molar relatively large, noticeably
constricted posteriorly (V.H.).

Description

The brown bear is a massive, very heavy bodied animal. In all its
parts, it is quite proportioned, although at first glance, it appears very
“clumsy”. This impression is also due to the fact that movement of
the animal is usually slow and unhurried. Posterior part of body—
croup and thigh—more massive than anterior; however, fore limbs
large, strong, and no such disproportionality between anterior and
posterior parts of body as in polar bear. Above-mentioned unconformity
is concealed in that withers are high, and its hair very long, giving
the impression of a sort of “hump”. This is also emphasized in that
the bear carrys the head lowered and raises it only while listening and
sniffing.

Head large and heavy, with broad and high forehead and ears
quite tall and set apart. It is only moderately elongated, and seems
less elongate than black and polar bears, but is relatively larger, more
massive and heavier. Ears comparatively small, rounded, and in

410

605

animal in winter pelage, basal half hidden in fur of head. Eyes very
small, dark; bare part on tip of nose quite large, black. Lips black,
large, quite loose, and like nose, very mobile (protrusible). Tail very
short and entirely hidden in fur. Claws very large and curved; those
on fore-limbs considerably longer than on hind-limbs; length of larg-
est may attain 5—6, or even 7-10 cm along curve. Their color is
blackish-horn or dark with lighter tip; in some forms, light-horn over
entire extent, or on greater part. |

On lower surface of hind foot, there are 5 well developed, bare
digital pads; base of free parts of toes covered with hair. Ail remain-
ing surface of sole bare and itself represents metatarsal callosities
(pads) united into one large area. Borders between them denoted by
fold beginning at outer part of sole. On lower surface of forefoot,
digital pads and large transversal anterior callosity in anterior part of
metacarpal region well developed; posterior to this, nearer exterior
side of sole, lies a small rounded callosity. Additional, ill-defined,
small secondary posterior callosity occurs closer to interior side of
sole.

Winter fur, especially in northern animals, very thick and long.
Length of longest hairs in large forms reaches 11-12 cm (on the
withers). Comparatively, general dimensions of hairs thin, but fur
quite rough to the touch; hairs flexible with silky luster, and as a
whole, the coat is very beautiful. Pile* of skin ill-defined, and on
living animal, full winter coat appears to flow when animal moves
(walks).

Summer, i.e., transitional fur (molt is once per year) is much
shorter and sparser. Apparently, in very old and large bears, fur is
shorter in comparison to that of younger, smaller animals. Length and
density of fur varies geographically. Fur of southern bears—Cauca-
sian, and especially Trans-Caucasian and Middle Asian—sparser and
coarser, of northern [bears] denser, longer, and more silky. Fur of
bears of the Far East and Siberia is especially good, and among them,
particularly that of Irkutsk and Trans-Baikaliya.

Color brown, generally homogenous over whole body. “Usual
tone of winter coat of adult Middle Russian bear is as follows: whole
back covered by hairs of dark and dark-brownish tone admixed with
various amounts of black. On lower part of back and on thighs, tone
of fur is darker, more blackish. On shoulder region, color is, on the

*Directional “set” of hairs on skin—Sci. Ed.

606

contrary, often lighter, possessing a slight reddish tint as a result of
the development of this color on the very tips of the hairs. Ears dark
brown and their intense color sharply distinguishes them from back-
ground of lighter forehead and upper part of cheeks, which are of
more reddish-straw color. Between the eyes, fur becomes darker again,
and merges into dark-brown or blackish-chestnut. This intense tone
covers the whole nose, around eyes, and lower part of cheeks. Chin
is also dark; on neck (below), a weak admixture of rusty fur is
observed; belly is dark brown. Limbs are darker than back; the nearer
the end of the feet, the blacker the intensity of fur color becomes.
Feet are dark blackish-brown. Length of hair in region of middle back
is about 55 mm, in shoulder region and between shoulder blades,
about 85, and on posterior part of back—about 90; on thighs—85,
and in middle of forehead, about 32 mm.

Color of individual hairs in region of middle back is as follows:
their deep base is dark-ashy-gray-brownish; from this basal portion
extends a wide, intensively-brown zone, and the extreme hair tips are
rusty-brownish. This color of the hair tips is weakly developed in the
middle back, and hence the general tone of the back is darker than
the sides, where the rusty tips of the hair are wider, and protrude
more noticeably externally. On the lower part of the back, rusty tips
are completely absent, which modifies it to a dark color. The hairs of
the very short tail are dark-brownish-blackish” (Ognev, 1931).

Individual variations from this widespread type of coloration are
comparatively small and are expressed only as darkening, lightening
or reddening of the general type. In extreme cases, Middle Russian
animals have a very dark, nearly blackish-brown color; in rare cases,
almost black or, on the contrary, light reddish-brown.

In some regions, individual variation is greater, and very dark or
very light animals are found equally—in extreme cases, golden-light-
pale yellow, or nearly straw-yellow. In other regions the whole gamut
of colors shifts in the direction of lightening. Sexual differences in
color are absent. Color of young is variable and in one litter there
may be both dark and light individuals. Generally, they are lighter
than are adults. In many cases in young bears, two well developed
white spots, not sharply outlined, are found on the neck and nape.
Quite often, these spots form a light collar. This latter, in rare cases,
is retained in subadult animals and, as a rare exception, (more often
in Central Asia) even in adults. By the second year, young bears are

411

607

generally lighter in color than adults and old animals, and among
them, very light individuals are often found.

Geographic variation in color is quite strongly expressed in some
parts of the range. In separate areas, there is a prevailing tendency
towards variation of the usual color intensity to one or the other side.
Thus, bears of the Caucasus, and particularly the Trans-Caucasus and
Middle Asia, are lighter than northern ones; contrast in color of the
limbs and body changes, appearing to darken in the region of the
withers, etc.

Skull of brown bear large and massive, with well developed
crests and general sculpturing. Sagital crest in adult animals is devel-
oped on frontal bones, and at place it unites with occipital crests; a
large posteriorly directed projection is formed. Postorbital processes
large. Facial region large and strong; its length approximately equal
to that of brain case—distance from anterior point of premaxillae to
line uniting ends of postorbital processes equal or almost equal to
distance from above-mentioned line to posterior point of dorsal pro-
trusion of occipital and sagittal crests. Distance from anterior margin
of orbit to anterior internal point of premaxilla greater than width of
frontal area between outer points of postorbital processes.

Anterior part of rostral region comparatively wide—its width
above canine teeth usually not less or hardly less than width across
preorbital foramina. Frontal area wide, sharply demarcated posteri-
orly by anterior branches of sagittal crest, and flattened or concave
in longitudinal direction. Braincase relatively small, not swollen, and
elongated. Zygomatic arches strong, widely separated. Mastoid pro-
cesses very large, articular [processes] massive and large. Nasal
opening relatively large and wide. Lower jaw massive, elongated,
with large coronoid process and very massive articular condyle.

Individual variation of skull very great. Skull may be relatively
narrower and more elongated, or broad with stronger or less pro-
nounced crests and general sculpturing; form, dimensions, and surface
(concave, flattened) of frontal area, size of postorbital processes,
steepness of transition from nasal region to the forehead, entire upper
line of profile, and a series of other characters may vary. Particularly
characteristic of adult animals is that skulls may be of two types.
One—“normal’—is large, with sharply defined protuberances, crests,
etc., and with compressed brain case; the other is of smaller dimen-
sions, with more weakly defined protuberances, crests, etc., and a
relatively less compressed and more swollen braincase; on the whole

608

a more “infantile” type. Such skulls are encountered, apparently,
throughout the whole range, but more often in several places therein
(see below).

Sexual differences in structure of the skull are well pronounced.
In the female, the skull is smaller and not so massive, crests and
protuberances of the skull are weakly developed, braincase is rela-
tively less compressed, zygomata not so massive and not so widely
separated, and the nasal opening is relatively narrow.

Age variation in the skull is very great and lies in relative reduc-
tion of the braincase, increase of facial part, development of crests
and processes and widening of the zygomatic arches.

Geographic variation of the skull is quite significant, but reveals
itself chiefly in general dimensions. Structural differences are more
weakly developed and express themselves mainly in the degree of
strengthening of protuberances, crests, etc., of the skull; in the lesser
or greater elevation of the frontal region, general relative elongation

Fig. 150. Upper [cheek] teeth of brown bear, Ursus arctos L. (left), polar bear,
Ursus maritimus Phips*, and white-chested bear, Ursus thibetanus G. Cuv.
Sketch by N.N. Kondakov.

*Sic; should be Phipps—Sci. Ed.

413

609

of the skull, and in width of the zygomatic arches. The relative
number of individuals in a population possessing the described “in-
fantile” skull is also significant.

Teeth very strong. Incisors relatively big, canines large, with
lower ones strongly curved backwards. First three premolars of
upper jaw underdeveloped, single-crowned, with one root. Usually,
first of these teeth lying adjacent to canine retained with age, as
well as third, shifted towards fourth premolar. Second, which is
smaller than first and third, very often absent in adult animals. It
usually falls out early, leaving no trace of its alveolus in the jaw.
More rarely, first or third or even both fall out and their alveoli
grow over. Fourth premolar much larger than others, with two roots
and never falls out—rudiments of premolars, entirely characteristi-
cally for family in general, do not encroach on it.

Second molar very large—broad and elongated. Its length ap-
proximately equal or slightly less, sometimes even more than, length
of first molar and fourth premolar together. It is approximately
twice as large as first molar, and length of both molars usually more
than width of palate between first molars (Fig. 150).

In lower jaw, first three premolars also very weak. Second one
absent in majority of cases, its alveolus usually not evident, first and
third also often fall out. Fourth premolar* always present and has
two roots; the first molar elongated and relatively narrow, second
approximately equal to it in length, but wider and more massive. It
is largest molar. Third molar shorter than the second, in anterior
part it has same width, posteriorly, somewhat narrower than [sec-
ond]; it [third] has rounded-oval outline; anteriorly, somewhat wider;
ratio of length to width approximately | : 1.5.

Brown bear, equally with polar bear, are the largest species of
Recent terrestrial carnivores. Its dimensions fluctuate very greatly
according to sex, age, individual and geographic variation, and also
by season. Moreover, accurate data concerning size of the bears
are quite scarce, with many contradictions. Therefore, character-
ization of the species in this respect is difficult. Body length of old
males of the largest races (Far Eastern, northwestern North
American) reaches 245—255 cm, height at withers—120—135 cm.
Weight reaches 500, 525, and 640 kg. A large bear, standing on the
hind feet, has a “stature” of 2.73—3.00 т.

*Misprinted “molar” in original Russian—Sci. Ed.

610

‘>
=.
Si

И
he
Sy

GL 17
и.’ “My

Е:

TE,
я
veh

Wit

412 Fig. 151. Skull of the brown bear, Ursus (Ursus) arctos L.

414

611

Such huge animals significantly exceed the average weight level
of their populations. They are usually very old animals, growing and
surviving under very favorable conditions (for example, preserves)
and particularly strongly fattened in winter. The weight of “nor-
mal” large males, as a rule, does not exceed 250—300 kg, or
somewhat more (320 kg—20 puds). Higher weight in Europe and
Asia (except the Far East) constitute rare exceptions, and in the
Far East are met with very rarely’.

The smallest races of bears have, apparently, a body length of
130—150 cm, and a weight of about 56—80 kg.

Variation in dimensions of animals in one or another population
depend to a considerable extent on the intensity of hunting in a
given region. Bears grow for a long time, and where they are much
hunted, the animals do not grow to maximum size, and in such
population, the proportion of young small bears is relatively great.
This occurs locally, for example, in the European part of the Soviet
Union. Large dimensions of Siberian and Far Eastern bears are
explained, not only by natural geographic variation of the species,
but in part evidently by the above-mentioned situation.

‘Olt must be taken into consideration that animal weight is often exaggerated
as hunters usually estimate it “by eye’—-very rarely is it possible to get accurate
weight of a dead animal. They are usually not very interested because they do not
deal with this account of the trophy. According to recollections of our famous
hunters and sportsmen of the end of the past and beginning of the present century,
S.A. Buturlin, G.I. Polyakov and F.F. Shillinger, a weight of 500 kg (30 puds) is
very rare (V.G. Heptner).

Records exist (Ognev, 1931; Couturier, 1954) of bears weighing 640 kg (40
puds, Kamchatka) and' 550, 589, and 750 kg (Alaska). Finally, one animal from
Kodiak Island, living in the Berlin Zoological Garden in 1937 had an unbelievable
weight—1200 kg. If all of these figures are true, then they belong to giants, which
occasionally are encountered among many species of mammals.

Very often, not only in popular, but also scientific literature, measurements
of North American bears are exaggerated, especially from the northwest where the
largest forms of the New World actually live. According to the most recent data,
“large grizzlies” weigh 250—300 kg (Couturier, 1954); animals from Alaska (very
large form middendorffii) have a body length of 180—190 cm (Bee and Hall, 1956);
according to Anthony (1928), body length of American bears is 180-255 cm,
shoulder height is from 90 to nearly 120 cm, weights from 158 to 410 kg and in
“park” animals up to 525 kg. Only Burt and Grossenheider (1952), followed
apparently by Trautman (1963), indicate body length of the form middendorffii
as about 240 cm., shoulder height 120—135 cm and weight to 681 kg (1,500 English
pounds) and more. Our Far Eastern bears correspond in their dimensions to the
large American races (see section “Geographic Variation’).

612

Sexual differences in measurements are pronounced—females
are always considerably smaller than males. Their average weight
consists, apparently, of not more than 75% of that of the males, and
evidently usually less (in the Pyrenees, the largest known weight of
a female bear is 250 kg, of a male—350 kg; Couturier, 1954).

Weight of animals changes greatly by season. Lowest weight
of all is observed in spring after winter hibernation, and in autumn,
in a well-fed animal, fat may constitute 20% and more of total
weight. Especially fat bears occur along the shores of the Pacific
Ocean, fattening on salmonid fishes. Age changes in measurements
and weights of bears are extraordinarily great. This is partially
explained by the fact that young are born very small and have a
weight of 250—400 g (Europe), or 450—625 g (large American forms;
Couturier, 1954). This constitutes approximately 1/500, 1/600 of the
weight of adult animals.

Dimensions of the skull of bears living in our country (excluding
subspecies differences) are as follows: greatest skull length of adult
males, 311-455 mm; of females, 275—397 mm; condylobasal length
of skull of males, 261—418 mm; of females, 258—373 mm; zygomatic
width of males, 175—277 mm; and of females, 147—247 mm. Sexual
variations in skull dimensions are relatively less than in general
[body] measurements (V.H.).

Systematic Position

Among some features of skull structure, the brown bear manifests
characters of quite high specialization. Among those are attested
the relatively elongated facial region, comparatively small brain case,
strongly developed crests, most of all the sagittal, and general sculp-
turing of the skull. In this regard, the subgenus Selenarctos
(white-chested bear, U. thibetanus) has less specialized charac-
ters. The shortened facial region, relatively voluminous brain case,
less developed crests, and several other characters, give the skull
of the white-chested bear a more infantile appearance in a sense.

At the same time, some features of body structure, for ex-
ample, the relationship of fore and hind limb measurements,
development of the proximal parts of the feet, and several others,
allow us to see in the [Asian] black bear a more specialized climb-
ing form than in the almost purely terrestrial brown bear. It is more
specialized as an herbivorous form. The above-indicated characters

416

613

of skull structure may be associated in part with the generally
smaller dimensions of the animal. These statements allow us to
consider the brown bear as qualitatively the “central”, more gen-
eralized form of the genus''. The polar bear itself is considered to
be derived from the brown, and in any event, stands closer to it that
the white-chested bear (V.H.).

Geographic Distribution

Eurasia, on the north to the limits of woody vegetation, on the south
to the Himalayas, Mediterranean Sea, and northwestern Africa,
North America west of longitude 90° W. Long., in north nearly to
northern extremity of continent, and south to Mexico.

Geographic Range in the Soviet Union

Range (reconstructed) occupies the entire forest, and in part the
forest-steppe and steppe zones, eastern tundras, the Caucasus and
montane regions of Middle Asia. It constitutes a considerable part
of the range of the species (about half), and occupies the greater
part of the territory of the USSR.

Within the USSR, the range consists of three, more or less
isolated, principal sections—the main European—-Siberian massif,
associated with forest and in part, forest-steppe and steppe; the
Caucasian, mainly the montane forest region; and the Middle Asian
part, where bears inhabit mountains that are partially unforested.
All three sections join, or were joined in the past, in the south,
beyond the borders of our country; the Caucasian with the Euro-
pean-Siberian through Asia Minor, the Middle Asian with the other
two through Iran, Afghanistan and China. During the historical
period, the range of the bear in our country changed greatly as a
result of its reduction. In previous times, there was less isolation of
the separate parts of the range in our [country], including the
European-Siberian and the Middle Asian parts. Evidently, such
contact occurred to the east of Kazakhstan, and in ancient times
also through the western Cis-Caucasus.

"This conclusion is preliminary. It is probable that the American black bear
must be considered the “central form”. The solution to this question is possible
only through a complete revision of all bears, the system of which, as stated,
cannot be considered accurate.

‘Seq-jodoy pure ч$Ате 1, ‘зиетуе4лео) ul ээиэл90
JO зио1Зэл рэуе[оз-—Чуп0$ Ul :втрипз о} $10155918зивц JO зи леу эзеотрит YOU ит змолту ‘359104 YSYZOAO[IG Ul 9949141590
Jo зито4 рэзе|оз-—ф ‘еыэат$ и1э5элм JO елрипз ози! SUOISSOISSULI) 121391 JO 1эр104 изэчом-—© 0961 иг рие Аицизэ Чэциэлмз 94}
JO $06 Jo pua зе 4551 Jo ued ивэдолая pure е11э91$ и123$9м Ul UOHINGINSIP Jo 1эр104 194195 —С ‘эЗив1 рэ32п13$40994 JO JOpPIOgG—|]
:(1э434ЭэН '9°Л) USSN Ш! ‘1 501248 (51541) 57541] ‘1еэд UMOI JO osury ‘7S| ‘SI SIv

Ovt 021

ЕЕ

417

615

The range has changed significantly in the past ten years, and
is changing quite quickly before our eyes. For this reason, a more
or less precise determination of the distribution limit for the animal
is, in many cases, impossible. Moreover, in some regions in the
north, bears roam very widely and delimiting the area of normal
permanent occurrence, the region of regular transgressions, and the
region of rare, especially long transgressions, is difficult.

The northern border of bear distribution in the European-Sibe-
rian part of the range, generally speaking, is associated with the
northern border of the forest and forest-tundra. In it the bear is
rare, although in some parts it remains permanently, and regular
transgressions occur. Almost everywhere, the animals go into the
tundra, mainly its southern part, but in some places they penetrate
quite far north.

On the Kola Peninsula, the region of permanent occurrence of
the animal does not reach the Murmansk coast. The border passes
along the latitude of Murmansk and extends to the mouth of the
Ponoi on the eastern shore of the peninsula. Wandering animals go
into the tundra in the summer time and to the north and east they
reach the seacoast. On the Solovets Islands, the bear is absent, and
probably was never present. Farther to the east, the border includes
the extreme lower reaches and the mouth of the Mezen’, the south-
ernmost part of the Kanin (mainly transgressions) and goes along
the southern border and southern extreme of the Timansk,
Malozeml’sk and Bol’shezemel’sk tundras, reaching the extreme
lower Pechora. In the area between the Kanin and Pechora, bears
in summer go far into the tundra and even reach the seacoast.

In the northern Urals, the bear normally lives below 65° and is
met with in the tundra up to 67° N. Lat. Farther east the border
goes to the extreme lower reaches and the mouth of the Ob’,
passing it at approximately the latitude of the Arctic Circle at
Salekhard. From here, the border takes its direction along the south-
erm shore of Obsk Bay or slightly before it. In any event, in summer
time the animals live along the coast itself. Farther to the east, the
border of the area of permanent occurrence is an arc-shaped line,
slightly ascending to the north, going to the mouth of the Taz on
Tazovsk Bay. In the Taz-Yenisei interfluve, the border ascends still
farther to the north, including the Great and Small Khetta [rivers],
and goes on to the Yenisei at Dudinka (69°30’ М. Lat.).

616

In the expanse between the northern Urals and the Yenisei, the
northern border of the area of irregular occurrence and trangressions
crosses the sources of the Usa (about 68° N. Lat.), somewhat
north of Yarro-to Lake and across Cape Kamenny (about 68°30’
N. Lat.) on Yamal, along the northern shore of the Tazovsk Pen-
insula (about 69° N. Lat.), across the head of the Gyda river on
Gydansk Peninsula (70°30’ М. Lat.) and ascends on the left [west]
bank of the Yenisei to the level of Tolstyi Nos [cape] (70°15’ М.
Lat.) and even further north (Shirokaya Bay). The outlined area
occupies not only the whole forest-tundra, but also a considerable
part of the southern tundra.

From the mouth of Yenisei, the border directs itself to the
mouth of the Khatanga, covering the basin of Pyasinsk Lake (69°30°
N. Lat.), the basin of the Khetta and the left tributary of the
Dudypta (Kamennaya) to 71°30’ М. Lat. and extends to the
Khatanga at 72°30’ М. Lat. (Novaya river). Thus, on Taimyr, the
area of more or less regular occurrence and short transgressions
(their delineation is difficult) goes farthest to the north and includes
true tundra. Here, their migrations also extend farthest to the north.

Farther to the east, the border is very poorly elucidated. It is
possible to consider that it goes along the extreme lower reaches
of the Olenek, Lena, Omolon, Yana, Indigirka and Kolyma, and on
the latter, proceeds to the mouth. On the Lena, bear dens, though
rare, occur at Виши (70°30’) and Kumakh-Surt (71°30) not far
from the beginning of the delta. Transgressions of the bears occur
northward—at Bykovsk Cape (72° N. Lat.).

In all the above-mentioned extent of Middle and in part East
Siberia, the border of the range generally proceeds along the border
of the krummholz and tundra, so that the krummholz also serves as
the region of the normal occurrence of the animal, while the area
of tundra is mainly that of summer transgressions”.

Eastward from the Kolyma, the border of the range goes con-
siderably to the north of the tundra border, proceeding along the
northern slope of the Anadyr range and extending to the Pacific

'?The border of the range in the European part of the Union and in Siberia
is mainly from Middendorf, 1867; Pleske, 1887; Flerov, 1929, 1933; Yakovlev,
1930; N. Naumov, 1930; S. Naumov, 1931; Ognev, 1931; Tugarinov, Smirnov
and Ivanov, 1934; Adlerberg, 1935; Heptner, 1936; Kolyushev, 1936; Shvarts,
Pavlinin and Danilov, 1951; I. Laptev, 1958; Belyk, 1953; V.N. Skalon and other
sources.

418

617

Ocean somewhere in the middle part of the Chukotsk Peninsula
(north of Krest Gulf).

Summarizing the previous data, it is believed that the northern
border of normal habitat occupied by the bear goes (except for
northeastern Siberia) along the northern border of the forest. How-
ever, in summer, animals (mainly males, apparently) roam quite
widely and actually wander everywhere in the southern tundra, and
strictly speaking, the actual border is located in this zone. In addi-
tion to this regular occurrence there are, in places, much farther
transgressions of animals to the north for distances of tens and
even hundreds of kilometers. In addition to the above-mentioned
transgressions in the European part of the country and in western
Siberia, especially distant transgressions into Taimyr to 73° and
even to Taimyrsk Lake at 74°, and to the ocean in Yakutiya, are
known.

The shore of the Pacific Ocean, to the southern border of the
state, forms the eastern border of the range. Bears exist on
Karaginsk Island, Shumshu and Paramushir on the northern Kurils
and on Kunashir and Iturup of the southern [islands] (absent on
remainder), on Sakhalin and Shantar islands. The northern and
eastern borders of the range at the present time are the same as
described and apparently have not changed substantially in any
way in the last 100 years. In contrast, on Shumshu Island the bear
has recently disappeared (Podkovyrkin, 1960).

The southern (reconstructed) border of the range in our country
extends from the Pacific Ocean to the Altai and Tarbagatai, coin-
ciding with the state boundary. Animals are absent and never were
present, apparently, in the steppes of southeastern and probably,
southwestern Trans-Baikaliya. This border has not been subjected
to significant change during the past 100 years.

From the Tarbagatai [range], the border generally proceeds
toward the northwest, crossing the steppes of Kazakhstan to the
Urals. It envelops, from the south, the Kazakh melkosopochnik
[small hills] (Kazakh folded country), passing somewhere in the
middle between Karkaralinsk and the northern shore of [Lake]
Balkhash. Farther, crossing the Nura and the upper Ishim, the
border encloses, from the south, the Kokchetavsk upland and passes
westward to the upper Tobol, crossing it a little south of Kustanai
(Ara-Karagai pine forest). Thence, the line of the border surrounds,
from the south, a section occupied by the animal in the former

618

Troitsk county (Kaban-Karagai), goes on to the upper Ayat (tribu-
tary of the Tobol entering above Kustanai), and thence to the valley
of the Ural, approximately at Orsk. The occurrence of the bear
here is, in particular, recorded at Rossypnaya and Nizhne-Ozernaya
(below Orenburg). The bear was distributed along the Ural valley
to Ural’sk.

In these demarcated steppe and forest-steppe districts of
Kazakhstan, the bear was distributed only sporadically. It lived in
pine forest islands, cut-over land and montane sections
(melkosopochnik) with woody and shrubby vegetation, and even
without it. The bear still inhabited some parts of Kazakhstan in the
18th century (sources of the Nura south of Karkaralinsk, Ara-
Karagai), and in other, it existed up to the beginning of the 20th
century (Sandyktau, Kokchetavsk mountains around Borovyi). It
cannot be excluded that in more ancient times the bear was distrib-
uted farther south than is now known and described here. Thus, it
is not excluded that it inhabited Ulutau, although there is no infor-
mation on this, and several other places to the south. It is interesting
that all of the border demarcated goes through the steppe zone, in
the east, very near the border of semidesert and desert zones.

The area inhabited by the bear in Kazakhstan itself represents
the southern extreme of the middle- and western Siberian part of
the range. However, in the steppe and forest steppe of western
Siberia the bear has not been present for a long time, and it is
difficult to explain how it disappeared in this region. Apparently, in
Kazakhstan, the bear still existed at some points even when it did
not exist in considerable areas farther north.

The recent (1950’s) the line of the southern border of the range
in western Siberia cannot be delimited with sufficient accuracy due
to inadequate information. In the Altai, the animals live everywhere,
including the extreme south—the basin of Markakol’ Lake and to
the Zaisan depression (in the depression itself, it is absent), in
Narymsk range and in Kalbinsk Altai (left bank of Irtysh). Farther,
it goes along the border of the more or less high trunked forest, 1.е.
along the northern foothills of the Altai, and, enveloping this mon-
tane country in an arc, surrounds the Kuznetsk Alatau from the
west and north, going on to Tomsk and thence to Novosibirsk.
Spreading over the Barabinsk steppe from the north and proceeding
slightly to the north of Lake Chana, the border of the range crosses
the Irtysh a little south of 56° N. Lat., and the Ishim—somewhat

419

619

north of this degree, going on to Tyumen’ and approaching the Ural
range, covering Sverdlovsk region a little north and west (45—50
km) of the city. Along Urals, the range of the bear descends to the
south as a large cape, reaching to 52° N. Lat., in the east including
the sources of the Ural [river]'*. Thus, in western Siberia, the
contemporary range of the bear already does not occupy the steppe
and the forest-steppe, and its southern border proceeds along the

‘southern part of the forest (taiga) zone. In some sections of Siberia,

for example in Cis-Baikaliya, in some places inside the range con-
siderable areas have formed where the bear had been a
comparatively short time earlier a common species, but has com-
pletely or almost completely disappeared. This was, unfortunately,
favored by the attitude toward the bear as a predator, the hunting
of which was in no way restricted.

In the European part of the Union, the natural range of the bear
in the south occupied not only the entire southern part of the modern
forest zone, but also the forest-steppe belt and was spread far
within the limits of the modern steppe".

Between the Volga and Ural, the animals lived along the Kinel’,
Samara, and in Zhiguli. The southern border, however, lay to the
south. Passing from the Ural river (Ural’sk), it apparently included
Obshchii Syrt and its southern spurs, extended to the Irgiz, and
along it to the Volga. Thus, in this area, animals were also suffi-
ciently widely distributed in the steppe zone. In some places, bears
apparently moved farther to the south than has been stated.

The distribution of the animal in the Volga valley is not clear,

13 Border of range from Tarbagatai to the Urals according to Ognev, 1931;
Shvarts, Pavlinin and Danilov, 1951; Kirikov, 1952, 1959; Yakushevich and
Blagoveshchenskii, 1952; Sludskii, 1953; I.P. Laptev, 1958; Afanas’ev, 1960 and
other sources.

№ In ancient times, our southern steppes were much richer in forests than in
later times. The southern limit of forests from the Carpathians and Moldavia
passed, apparently, approximately along 48° N. Lat., through the upper Ingul,
Dnepr, the upper Berda and Mius to the lower Donets. To a considerable degree,
these regions were bottom-land forests. Along the Dnepr, the bottom-land in the
form of quite considerable massifs reached even to its mouth, and the same thing
took place along the Bug and along other southern rivers. Considerable woody
vegetation existed along the Don and its tributaries. The bear is ecologically very
flexible and could have existed not only in the presence of entirely insignificant
forest vegetation as in Kazakhstan, but also along the bottom-land forests, in
shrublands along the river valleys, and even in the steppe.

620

but probably they extended along the bottom-lands quite far to the
south since they were, apparently, quite widely distributed in the
forest-steppe and steppe zone in the Don basin. Here, the animals
lived along the entire Medveditsa to its mouth, and along its tribu-
taries the Ters, Knyazevk, Karamysh, Idolga and Kamyshlei. Along
the Don itself, bears lived not only in the north (Shipov forest
around Pavlovsk) but also considerably lower—to Starogrigor’evsk
station, at the mouths of the Khopr and Medveditsa, and even in the
Kletskaya region. Along the Donets, bears lived mainly in the north—
at Chuguev, Zmiev and in other places to the south, southeast and
southwest of Khar’kov, but even in the region of the Oskol mouth,
and they reached the location of present-day Lugansk.

To the west, the animals were distributed in Chernigov and
Kiev districts and near Poltava. More accurate distribution of the
animal along the left bank of the Dnepr is not known; however, it
apparently descended to the south considerably beyond Poltava. It
may be assumed that the border of the range went from the Donets
to the Dnepr approximately at Zaporozh’e. To the south on the right
bank of the Dnepr, there is some information concerning the occur-
rence of bears in Chermyi forest near Kirovograd, in the Savransk
steppe southeast of Balta at the mouth of the Samotkan’ river, and
even in the steppes around Ochakov, on the lower Dnepr and at
Perekop, 1.е. bears were distributed to the shore of the Black Sea.'°

Therefore, in the European part of the Union, bears were dis-
tributed not only in the forest-steppe but were widely encountered
in the steppe zone, in particular, apparently, in its western part."

In the west, the range of the bear in the past extended to the
Baltic Sea and to the southwestern foot of the Carpathians—to the
Pannonian plain.

Information concerning Ural’sk, Volga, Don basin and Chernyi for-
est is from the 18th c[entury]; data about Balta, lower Dnepr, Ochakov
and Perekop, to the 16th—17th [centuries]. Fossil remains of bears
from the Pleistocene are known from the Crimea.

'5Вогдег of range in European part of Union according to Kirikov, 1952, 1955,
1959; in part Korneev, 1952; Sokur, 1960 and other sources.

‘SOchakov bears were even called by the author who described them
(Bronevskii) “steppe bears” “Ursi campestres” (cited from Kirikov, 1952). In
this respect, our southern bears (as in Kazakhstan) are entirely analogous to some
American forms.

420

621

The described reconstructed southern limit of the range of bears
in the European part of the Union, in contrast to the northern, has
been very strongly changed during the last century, and displaced
northward for hundreds of kilometers—in places almost 1000 kilo-
meters and perhaps more. An entirely intensive process of range
reduction has gone on in the 20th century, particularly in the last ten
years. The border did not change so quickly during the entire [pre-
vious] history of the species. Not only the deficiency in the accurate
data, but also to a less degree, the stated situation does not make
it possible to give the contemporary southern border of the range
with sufficient accuracy—it changes before the eyes from year to
year. It has changed more and, apparently, more quickly, than the
southern limit in Siberia.

In the 40’s and 50’s of our century (until 1960), the southern
limit of distribution of the bear in the Ural [mountains] and the
European part of the country might be outlined as follows. Starting
at the eastern slope of the Ural, approximately 50 km west of
Sverdlovsk, it passes directly southward along the eastern edge of
the Ural forests, occupying the range (Shaitantau) south of approxi-
mately 52° N. Lat. Thence, the border turns sharply northward,
defining a Ural “extension” of the range on the west. The border
goes along the foothills west of the Ural without crossing the Beluya
[river]. Somewhere in the vicinity of 60° N. Lat. it curves sharply
to the west, dividing the northeastern part of Perm district, where
bears still exist, from the remainder, in which the animal is already
extinct. The westward trend of the border quickly changes to the
southwest and then to western again—the border, descending to the
lower Kama, bypasses it from the north and crosses the lower
reaches of Vyatka, thus encircling Tatar Republic from the north.
Here the bear, as a permanent inhabitant, was already absent at the
end of the 20’s—-start of the 30’s, although individual transgressions
moving down from the northwest (from the Marii ASSR) still oc-
curred locally in the 40’s, 50’s, and even 60s. Migrating animals
almost immediately fall prey [to hunters] (V.A. Popov).

Crossing the Volga, apparently somewhere in the vicinity of 48°
E. Long., the border goes on steeply southward, extending over the
forests of Sura basin and descending here quite far to the south,
approximately to 54° N. Lat. Thence, turning somewhat northward,
the border proceeds along the level of Temnikov in the Mordovsk
Republic, westwards to the Moksha, and through the forests of the

622

Tsna basin, producing a long, very narrow, extension southward.
However, this extension does not reach Tambov. These data be-
long to the past decades; however in 1960, the bear was already
absent throughout the forests of Tsna and Moksha except the
Sarovsk forests (Zametchinskii region, K.I. Nagornov).

From the region around the mouth of the Moksha, the border
proceeds somewhere along the left bank of the Oka [river] at a
distance from it, then suddenly descends southward, making a loop
which includes Ryazansk Meshchera. The southern border of this
loop passes along the Pra river. From the Pra, the border turns
abruptly northward, and passing first through the regions of Vladimir
district, surrounds the Moscow [district] from the east, north and
west.

In the Moscow district, bears even a short time ago were quite
widely distributed and lived not far from Moscow. In 1891, a bear
started from a wolf’s den was killed at Pushkin (about 30 km from
Moscow along the Northern railway). Even in the 20’s, the animals
lived regularly in the former Bogorodsk (Noginsk), Dimitrovsk and
Klinsk counties (the east, northeast and north of the district). In the
20’s, they were still found in the then Dolgolugovsk forest planta-
tion, which included the vast forests around Khot’kov, Sofrin and
Pushkin (along the Northern railway—between Moscow and
Zagorsk); they also existed in the forests around Zagorsk and far-
ther north. The bear began to disappear quickly in the 30’s, and in
the 40’s and 50’s, was no longer in the nature of a permanent
inhabitant of this region; it very rarely appeared as a transient
moving from the north along the Dubna or from Meshchera to the
forests around Shatura, i.e. from the east and southeast (Ognev,
1931; Formozov, 1947; V.G. Heptner). The last of such transients
occurred locally on 7 December 1960 in the exceptionally warm
winter, when a shatun* was killed in the forests at Lukhovits
(newspaper “Vechernyaya Moskva”, 24 December 1960).

A somewhat exact location of the border in the Ivanovsk,
Yaroslavsk and Kalininsk district is impossible. Apparently, the range
includes the western parts of Orlovsk district and, perhaps, Kaluga
district, since in the 50’s it included all of Bryansk district except

421 its extreme southern parts (Brasovsk, Sevsk, Novozybkovsk,

*Starving, non-hibernating bear in winter; see p. 676—Sci. Ed.

623

Klimovsk, Klintsovsk and Starodubsk leskhoz [forest cooperatives];
(Fedosov and Nikitin, 1951).*

In Vladimir district the bear was already absent in the 50’s
(N.D. Sysoev).

In Byelorussia, where the bear was, a short time ago, widely
distributed almost everywhere, in 1950—1951 it was already found
only in the northern regions. The border passed farther north of
Mogilev and northeast of Minsk, but then turns sharply to the north-
west and north, and passes through the regions of Borisov,
Pleshchentis, Begoml’, Donshchitsa, Glubokov and Markovshchina.
A small center of occurrence of the animal, separated from the
Northern Byelorussian, is found in Belovezhsk Forest (Serzhanin,
1955). Farther to the north the limit occupies, apparently, the west-
ern extreme of Latvia (in Lithuania, the bear is now absent), going
into Estonia. Here, in 1976, the bear was quite widely distributed,
being encountered west of a line from Pyarnu (Riga Gulf)}—Cape
Yumind (Gulf of Finland), and in the north reaching to the Gulf of
Finland and in the south to 58° N. Lat. A separate section of
occurrence, which a short time ago was still connected with the
main part, is found northeast of Khapsala (Kh.I. Ling).

From northern Estonia, not so far from the coast of the Gulf of
Finland, the boundary passes eastwards to Leningrad. In the west
of Leningrad district, the range enters the regions of Luga, Slantsev
and Kingissen. Bypassing Leningrad to the south and east, the
border passes through Gatchina (about 40 km from Leningrad),
Vyritsa (60 km), Tosno (about 60 km), Lisino (45—50 km, along the
highway to Moscow) and Mga (about 40 km; N.K. Vereshchagin),
and goes on to the shore of Lake Ladoga and, surrounding it,
passes into Karelia. On the Karelian isthmus, the animal is absent.
The state boundary constitutes the western border of the range in
Karelia and farther north.

Inside the region defined by the above mentioned line of the
southern boundary, the distribution of the animal is very uneven
and, equally with regions where the population is found in a more
or less normal condition, there are considerable expanses where it
is reduced (western and eastern parts of Leningrad district), or the
animals are very rare or appear only as transients. In considerable

*No initial parenthesis in Russian original—Sci. Ed.

423

624

areas inside the range the bear is already completely absent. It is
impossible to characterize various parts of the range in this respect,
not only because of the absence of information in the literature, but
also because the distribution and numbers of bears changes very
quickly in a negative direction—literally in individual years. Thus, if
in the beginning of the century the bear was common at any place
in the northern districts of the Ukraine, by the end of the forties,
only individual transient animals from Byelorussia to the northern
parts of Chernigov, Zhitomirsk and Volynsk districts were met with,
and later, they completely disappeared. In Byelorussia a short time
ago, the animal was distributed almost everywhere, while at the
present time, its range is restricted го the above-described northern
part, and so on.

The picture which is shown of changes in distribution and, in
part, numbers of bears at the southern limit of the contemporary
range, are indicated by special investigations carried out in 24 middle
districts in 1962. These districts lie from the upper Dnepr in the
west (boundary of Byelorussia) to the Kama in the east. Within this
territory, the bear inhabits only forest massifs of large area, there-
fore the southern limit of its distribution coincides quite exactly
with the recent limit of large expanses of forest. In Smolensk
district, the border passes along the right bank of the Dnepr, and
in the neighborhood of Dorogobuzh, a small amount of bears are
encountered in forests on the left bank of the Dnepr, which are rich
in swamps. From Dorogobuzh, the boundary ascends north-north-
east as far as the village of Sychevka, whence it passes along the
left bank of Volga to Kalinin and beyond to Bezhetsk and eastward
to Uglich and Tutaev. Farther on, the boundary again goes long the
right bank of the Volga to Zelenodol’sk (Tatariya), thence ascends
northeast towards to Malmyzh, Vyatskie Polyana and Izhevsk.

North of this line during the time indicated, the bear is met with
everywhere; however, to the south, there were only a few separate
sections of its occurrence:

1. the forest massif between the cities of Kostyukevichi, Kletnya
and Roslavl’ within Bryansk and Smolensk districts (up to 10—15
individuals);

2. the forest massif along the left bank of Desna, in the inter-
fluve between the Zhizdr and Resset rivers, and the sources of
Vytebet’ river within Bryansk, Kaluga and Orlov districts (from 20
to 30 animals);

|
}

Ы.]
\
o

Fig. 153. Details of the distribution of the brown bear Ursus (Ursus)
arctos L. in 24 districts of Middle Russia (outlined by dashed line—3)
from the upper Dnepr (eastern boundary of Byelorussia) to the Kama.
In the north (1) district of more or less general occurrence. Outlined
separate sections (2) farther south of the general boundary—individual

422

separated from the general range in the north.

Data for 1962 (V.P. Teplov and S.G. Priklonski1).

isolated areas of occurrence

626

3. Meshchera forest massif along the left bank of the Oka in
Ryazan district (5—10 animals);

4. the forest massif along the banks of the Tsna, middle course
of the Moksha and right bank of the Oka within Tambovsk, Penzen,
Ryazan, and Gor’ki districts and Mordovian Republic (30—40 bears);

5. the forest massif along the right bank of the Sura in the
Chuvash Republic (15—20 animals);

6. the forest on the right bank of Kuibyshev reservoir south of
the city of Tetyusha (in 1960, a female with cub) [Fig. 153]."”

In the sixties, a transient bear was recorded in Taldomsk region
of Moscow district and near Vyazniki of Vladimir district (V.P.
Teplov and S.G. Priklonski1).

In 1960, as a remnant of their [once] wider range in the Eu-
ropean part of the Union, there is quite large, entirely isolated area
of bear habitat within our country in the Carpathians. It consists of
a small belt extending in a northwestern direction from the Roma-
nian border (from a place southwest of Chernovtsy) and nearly to
the state boundary with Poland, south of Drogobych. It is a region
of taller, denser Carpathian forests within Zakarpatsk [Trans-
Carpathia], Chernovitssk and L’vov districts. In particular, the range
includes Chernogor Gorgan, Beskid and Borzhavsk polonina* and
other parts. This section represents a continuation of the range of
the bear in Romania."

The Caucasian section of the range in our country was, appar-
ently, not connected with the European-Siberian part during the
historical period. The connection was affected through Central
Europe, the Balkans and Asia Minor. However, in the Holocene
the northern population of bears was connected with the Caucasian
through the steppe region. Bears undoubtedly lived along the
bottom-lands of the southern Russian rivers and the rivers of the
northwestern Cis-Caucasus, and in steppe ravines and reeds of this
region. In the lower Don, the animals were encountered even in the

Information concerning occurrence of bears in Tatariya, except as previ-
ously mentioned, is considered unreliable (V.A. Popov).

* Local word for montane meadow—Sci. Ed.

'8 Described southern limit according to Ognev, 1931; Popov and Lukin, 1949;
Heptner, Morozova and Tsalkin, 1950; Kuklin, 1951; Fedosov and Nikitin, 1951;
Shvarts, Pavlinin and Danilov, 1951; Kirikov, 1952, 1959; Korneev, 1953; Serzhanin,
1955; Tatarinov, 1955; Savinov and Lobanov, 1958; Sokur, 1960 and other sources
and materials of N.K. Vereshchagin and V.G. Heptner.

425

627

8th—13th c[enturies]. The former direct connection between Cau-
casian and Russian bears is shown by several peculiarities of
geographic variation of bears of the Caucasus (see below).

In the past, the range covered all of the forest regions of the
country; i.e., in the main, the entire Caucasus—Great and Lesser, as
well as the western Trans-Caucasus from great heights to sea level.’
The animal was absent only in the steppe region of the eastern
Trans-Caucasus, although they penetrated along the (ига! forests of
the Kura into forestless districts at least to the level of Alazan’;
perhaps they were also found along some other rivers. It is possible
that the bear was absent in some forestless desert places in the
mountains of inner Dagestan, in the desert expanses of the Erevan
depression and, perhaps, in several other relatively small sections.

Along the Glavnyi [Main] Caucasian range, the range began at
Anapa in the west, and in the east reached the shore of the Caspian
[Sea]—to the forests in the foothills of Dagestan and the forests
along the lower Samur river and mouth. It occupied the entire
forest of the northern premontane and foothill area as far as Maikop,
Pyatigor’ (Zheleznovodsk), Nal’chik, Ordzhonikidze (Vladikavkaz)
and Khasav’yurt. The forested areas of the Stavropol’ uplands
were even included in its range.’

At the present time, the range of the bear in Caucasus has
been reduced, in some place considerably. In part this even took
place in the past century, and in part, even mainly, in our century—
in the second quarter of it. The bear is absent in Pyatigor’e (here
it only rarely intrudes), stretching the border, in part in connection
with great clearings, from the northern foothills; the bear is now
absent at Maikop, it has nearly disappeared from Kabarda and
Balkariya, at Khasav’yurt, etc. The animal has disappeared from
many regions of the western Trans-Caucasus (Kolkhida); the re-
gion of its occurrence in Armenia has somewhat decreased. The
bear is nowhere present in the steppes of the Trans-Caucasus, it
is almost never encountered in Talysh, etc.

In the Trans-Caucasian steppes at Stavropol’, it disappeared
more than 100 years ago; along the Black Sea coast, the bears still

‘Bears are able to live in nearly forestless mountains, transgressing to entirely
forestless desert mountains, and, in summer, are found everywhere in high-montane
(alpine) meadows.

20However, Dinnik denies the presence of bear in “‘Stavropol’ governance”.

(лэщаэн “D'A) рэзеотри! jou эле $и0155185иещ [eNpIAIpUl Jo soov[q “sed зиеззтр oy) UT adUeLINDD0 1915504
JO suoISo1 0} 1ээл змеш иопзэпо “1 SojouD (515411) 5п54] ‘1еэ9 UMOIG OY) JO э8иел $91945 pajonAsUOdeY “pC ] 14 УС

PAR et
ЗК а

- ie i ww Kt A
cae
yy 7) <
| wy

<) © SS

629

occurred nearly to the sea, even in our day; at Khasav’yurt it lived
in the 90’s [1890’s] and 10’s [1910’s]. In the beginning of the 20’s,
it was found in the forests of the foothills (“Chernye Gory”) ac-
tually only some kilometers from Ordzhonikidz (Vladikavkaz). Almost
everywhere, especially in the Lesser Caucasus, the number of
animals has been reduced.”' Accurate data on the recent distribu-
tion of the bear in the Caucasus are very few. Generally, its range
is connected with its direct pursuit, and with its reduction as forests
steadily decrease.

In Middle Asia, the range of the bear is associated with moun-
tains, such that the bear here lives in some places where there is
only a sparse growth of woody junipers or pistachios, or shrubs in
ravines; in some places, it exists in the entirely treeless, even desert
mountains.

The reconstructed range in Middle Asia and Semirech’e occu-
pies the Saur, Tarabagatai, Dzhungarsk Alatau and the whole of
the Tien Shan system including the western ranges and Karatau.
Furthermore, its distribution includes all the ranges of Pamir-Alai
system westward to the western parts of the Turkestan, Gissar
(V.G. Heptner; Leviev, 1939) and Darvazsk ranges. In Kugitangtau
and Babatag, and in some other elevated interfluves between the
right tributaries of the Amu-Dar’ya, the bear was absent from time
immemorial, or at the very least for a long time. Apparently, the
range did not include the Nuratinsk mountains. In Turkmentiya, the
range occupied the Kopet-Dag; however, in Great Balkhan, in the
mountains on the right bank of the upper Tedzhen (Gyaz’-Gyadyk),
and in the mountains east of Kushka (Chengurek mountains), the
bear was absent (V.G. Нершег).

It is impossible to clarify the recent range of the bear in Middle
Asia in more detail due to insufficiency of information. As com-
pared to the above-described, it undoubtedly changed during the
past century and particularly during the last ten years—the general
area of the range decreased, and significant “gaps” appeared in it,
but apparently the animal is still encountered in the majority of the
mountain ranges where it previously existed. However, it is now

"Distribution of the bear in the Caucasus according to Dinnik, 1914; Satunin,
1915; Smirnov, 1916, Ognev, 1931; Heptner and Formozov, 1941; Vereshchagin,
1947, 1959; Dal’, 1954; Tembotov, 1960, and other references, and according to
materials of V.G. Heptner.

426

630

absent in Karatau, and disappeared, or is only met with very rarely
as a transient from Iran, in the Kopet-Dag (V.G. Heptner).

Geographic Range outside the Soviet Union

In Asia, the reconstructed range occupies the islands of Hokkaido,
the Korean peninsula and northeastern China, except its central
steppe areas; i.e. mainly the mountains of the Great Khingan, II’khuri-
Alin’, Lesser Khingan, the eastern Manchurian montane region,
Chanbaishan’, Kentei-Alin’, Laoelin and Chzhanguantsalin. In the
Mongolian Republic, its distribution occupies, in the north, the Khentei
ranges and the Cis-Hobsogol part of the country southwards ap-
proximately to 48°30° М. Lat., at the boundary with the USSR
along the left bank of the Selenga river. In China, in the extreme
west, the range occupies the eastern part of the Tien Shan region,
in Kashgariya and Dzhungartya, in particular the Borokhoro moun-
tains at Barkul’. Thence, the range extends as far as the Trans-Altai
Gobi, the Gobi Altai, and the southeastern part of the Mongolian
Altai in the M[ongolian] P[eople’s] R[epublic].*

Farther, the range includes all of Tibet, including Nan Shan, and
Tsaidam, the region south of the great bend of the Huang he
(Tsing-ling mountains northward) and, apparently, northern and
western Sichuan. The question of the occurrence of bears in the
eastern part of China to the east of the described limits remains
open. It is not excluded that bear never existed there; however,
civilization has eliminated all natural relationships in this region so
that this cannot be clarified.

To the south, the Central Asian region of bear occurrence
covers the Himalayas, including Nepal, Kumaon, Kashmir with the
Karakorum and northern Punjab, and Waziristan. This animal as
described does not, apparently, penetrate further to the south (into
Baluchistan). In the east, the animal does not get into Burma (in-
formation concerning its occurrence in the Shan region is unverified).

In Afghanistan, the range occupies the regions associated with
the Pamir (Badakhshan, Wakhan), and the Hindu Kush. Details are
not known. In Iran, the range occupies on the one hand the ex-
treme northern region—forested districts associated with [Mt.]
Elburz—and extends as a band from the Talysh to Kopet-Dag. On

*Isolated population also in extreme southern Gobi desert, along border with
China—Sci. Ed.

631

the other hand, it proceeds as a wide projection from northern Iran
to the southeast along the western and southwestern part of the
country through Luristan approximately to Shiraz. In central, south-
ern and eastern Iran, the described species is absent. The range
occupies all of Asia Minor (montane sections), and montane
districts of northern and western Syria, Lebanon and Palestine.

In Europe, beyond our borders, the reconstructed range occu-
pies substantially all of the continent to the extreme north, including,
in the west, England and Ireland; southwards, it reaches to the
shores of the Mediterranean Sea including the Apennine and
Pyrenees peninsulas. In Sicily, Corsica and Sardinia, the bear was
absent as well as on the islands of the eastern part of the Sea.

In Africa, the bear occupied the Atlas mountains.

In North America, the range of the brown bear (reconstructed)
occupies the western half of the continent, approximately west of
90° W. Long. The southern limits of the range, starting along the
shore of the Pacific Ocean, include the northern part of the [Baja]
California peninsula. On the continent, continuing at some distance
from shore of the Gulf of California, it descends along the Mexican
plateau to the southern part of the province [state] of Durango,
approximately at 20° N. Lat. This is the southernmost point of
occurrence of the animal in the New World. Thence, the border
outlining this southern extension on the east goes northward through
northeastern Mexico (Coahuila) into Texas (USA), leaving its west-
ern part within the range boundary. Extending farther in a northern
direction, the boundary leaves within the range Oklahoma to the
west and the greater part of Kansas, and goes directly northward
through the western edge of Iowa and Minnesota and through the
eastern part of Manitoba, passing to the shore of the Hudson Bay
at Churchill. Running farther north along the western shore of the
Bay, the boundary turns to the northwest somewhere around
Chesterfeld Inlet and reaches the northern coast of the continent.
Along it, and in places some distance from it, the boundary pro-
ceeds westward, transecting the shore of the Bering Sea and
covering the Alaska Peninsula and Kodiak Island, continues along
the coast of the Pacific Ocean southward to the previously de-
scribed locality in California. Not included in the range, apparently,
is that-western part of the territory lying to the north of Bristol Bay
in which are found the lower Yukon and Kuskokwim [rivers]. The
brown bear is known from St. Lawrence Island and Unimak in the

427

632

Bering Sea; on others it is absent. Brown bears are absent also,
apparently, on the Queen Charlotte Islands and Vancouver Island.

The recent range of the brown bear differs very strongly from
that as outlined, and comprises only the smaller part of it. At the
same time, the range has been divided into separate, comparatively
small, sections—often.very small—which are entirely separated from
one other. In the whole of the range as outlined, there remain only
some more or less big areas inhabited by the animal. The largest
of these is the European-Siberian area within the USSR; the Cen-
tral Asian is quite large; and a considerable area exists in North
America.

In Europe at the present time, bears remain in Norway, Swe-
den, and Finland; in the French and Spanish Pyrenees; in the
Cantabrian mountains in Spain; in the Italian Alps, the middle part
of the Apennines; in Poland, Czechoslovakia, Hungary, Romania,
Bulgaria, Albania and the European part of Turkey. They are most
common in Scandinavia, in parts of Bulgaria and Yugoslavia, and
especially in Romania. In the other countries mentioned, the ani-
mals are very scarce and in part are found only as individual
specimens. In Africa, the bear became completely extinct long ago.

In Asia, outside the borders of our country the bear has been
conserved only in Asia Minor (in places), northern Iraq and in the
above mentioned part of Iran. In Japan, it has been exterminated.
The range on the Korean peninsula and in the Mongolian Republic
has decreased and continues to decrease. That part of the range
occupying Tibet and adjoining places in the Himalayas, is still large.

In America, the range has decreased very greatly. The animal
is still widely distributed in Alaska, in northern and western Canada
(except Manitoba), but in the United States, it is substantially, found
only in the Rocky Mountains in the states of Montana, Idaho,
Wyoming and Colorado. An isolated section of habitation is found
far to the south on the Mexican plateau.” It is evident that farther
reduction in range and numbers of the animal is inevitable (V.H.).

Geographic Variation

Geographic variation in bears has always attracted attention,
especially in the first half of our century. At the same time, many

Recent distribution of bear in Europe and America according to Couturier
(1954).

633

errors and misrepresentations have been made in its study such as,
so it seems, have not been made in the investigation of variation of
any other species of carnivorous mammals. Bears were the object
to which was applied a unique concept of Merriam’s (1918), who
accepted 77 “species” and 9 races for only the North American
part of the range, the majority of which he himself described. This
astonishing scheme is at present encountered in the American lit-
erature (Miller and Kellog, 1955).¥ A special genus of brown bears
was even described. This applies not only to the American forms,
where the misrepresentations are especially great, but also to ani-
mals of the Old World, including those living with us. The situation
noted, if one sets aside simple lack of ability and knowledge, is
explained partly by the undirected theoretical positions of some
authors and by the lack of study of the species question in the
recent past.

The large nongeographic variation in the bear (see above) cre-
ates essential objective difficulties. Particularly in this respect,
individual differences are important. In some places (Caucasus)
dark and light, large and small animals of the same age may live
side to side. The existence of animals of different dimensions in one
place is especially remarkable.

Individual differences in the skull are especially great—not only
in dimensions, but also in protuberances, crests, etc. profile, general
massiveness and others. The degree of tooth wear may be very
different in animals of the same age, which may perhaps be asso-
ciated with characteristics of foods of animals in various localities,
etc. Lastly, intensity of hunting may affect general dimensions of
the animals, and belonging to a definite population, as well as their
cranial characteristics. In one locality, the animals may live to greater
age and size than in another, and these differences may appear to
be entirely real systematic ones. It is quite natural that with a small
collection of material and a mainly morphological concept of spe-
cies and races, there was formed an exaggerated representation of
geographic variation in our bears (Smirnov, 1916). For our territory,
Ognev (1936) even accepted 6 species of bear, as well as one of

23Only in the most recent time has the species unity of the North American
brown bears been clearly stated (Rausch, 1962) and interesting data were presented
concerning the clinal type of variation in several important characters (condylobasal
length, i.e., general dimensions).

428

634

them with 8 races. This arrangement was held for a long time,
although even in the middle 30’s, more sensible opinions were
expressed about geographic variation of bears.

In general the trends in the Old World in geographic variation
in bears and their corresponding races can be traced in several
main directions.

1. European-Siberian bears (group arctos). Size moderate, in
places small, dark color type predominates, light type relatively
rare. Claws dark.

Europe, Caucasus, Siberia except the east, northern part of the
M[ongolian] P[eople’s] R[epublic].”

2. Near Eastern bears (group syriacus). Size moderate and
small. Light color type predominates, dark is rarely encountered.
Claws light or near-light.

Part of Trans-Caucasus, Palestine, Lebanon, Syria, Iraq, Asia
Minor, Iran, Afghanistan, western Himalayas, Pamir-Alai and Tien
Shan mountains.

3. Far Eastern bears (group piscator). Size large and very
large. Dark and very dark colors predominate, light color type is
also met with but more rarely than in European-Siberian bears.
Claws dark.

Far East and in part eastern Siberia from our southeastern
border to Kamchatka and extreme northeastern Siberia.

4. Tibetan bears (group pruinosus). Size moderate. Fur is par-
ticularly long and shaggy; light and dark color types are encountered
approximately equally (intermediate color predominates). Often lighter
on the neck (“collar”). Claws light. Skull distinguished by relatively
flattened choanae (internal nares), arch-like curve of molar row,
larger teeth and several other characteristics.

Tibet and several contiguous countries.

The Tibetan bear group differs from other groups more than
they differ from each other, and many authors considered this
group as an independent species (Flerov, 1935; Allen, 1938). For
this there is, apparently, not sufficient foundation, although the Ti-
betan group, to a certain sense, may be contrasted with all others.

Тре question concerning the North African U. a. crowtheri is set to one side.
This is a “mythical animal” (Harper, 1945) which became extinct more than 100
years ago, of which no specimens are preserved in museums.

429

635

Within the boundaries of our country, all of the racial groups
are represented, except the Tibetan. The relationship of Tien Shan
bears to them (to “species” pruinosus) 1s without foundation.

It is difficult to form an opinion about the groups of races
existing in America for the reasons given above. Apparently, there
are two main ones: 1) a group of large and very large bears of
northwestern Alaska and the coastal regions of the Pacific Ocean
southward to Dixon Strait [Entrance] (about 50° N. Lat.), including
Kodiak Island (forms beringianus, middendorffii, gyas and sev-
eral others); and 2) races of the “grizzly” group occupying all
remaining part of the species range in America (Haltenorth and
Trense, 1956). The first group is, undoubtedly, the same as the Far
Eastern one of the Old World (group piscator); the second is,
apparently, analogous (parallel) to the European-Siberian (group
arctos) and Near Eastern (group syriacus). There is obviously no
sharply delineated group such as the Tibetan (pruinosus) in the
New World.

After Ognev (1931) had published his unsuccessful scheme of
geographic variation in bears, this question was dealt with by many
authors as regards bears of the Old World. Only few of them
(Adlerberg, 1935; 1935a; Flerov, 1935) used good collections and
were able to contribute something substantive. The majority of
them (see review by Couturier, 1954) did not have sufficient ma-
terial from our country and combined data from literature. Their
review contributed little of use and sometimes confused the picture
even more. There is not yet a true contemporary analysis of geo-
graphic variation. The data given below are preliminary and only
note more well-defined subspecies.»

European-Siberian bears—group arctos

1. Middle Russian bear. Ursus (Ursus) arctos arctos Linnaeus,
1758 (synonyms, all the above mentioned synonyms, except those
mentioned below for other races).

Couturier’s opinion (1954), wholly denying the existence of subspecies of
brown bear is only “а gesture of despair” and of course, cannot be accepted. The
presence of real races is not in doubt, and the question lies only in their number,
in the evaluation of populations with intermediate characters (“transitional”), in
nomenclature, and so on. While denying the presence of subspecies in bear,
Couturier accepts, however, the occurrence of numerous separate “populations”,
which, of course, does not help in clarifying the picture.

636

Size moderate.** Pelage of moderate length, density and soft-
ness. Color varies from very dark blackish-brown with rusty-gray
highlights, depending on lighter terminal parts of guard hairs, to light
straw-brown with translucent dark-reddish-brown underfur. Under-
fur is always dark. Limbs are darker than body; in light forms,
muzzle is darker than trunk, dark spot (area) on withers absent.
Collar is sometimes developed, but is narrow, not covering entire
base of neck, or not developed on nape. Occasional individuals of
extreme color are almost black with a weak admixture of brown
tone, or almost straw-yellow.

Details of coloration of the two main types are as follows.
Light type: general tone is brownish-straw with translucent dark
underfur. Base of guard hairs and underwool red-brown, dark.
Muzzle brownish-rusty. Lower lip and chin darker. Forehead and
side of head gradually lightening, imperceptibly grading into light
color of neck and trunk. Ears dark, same tone as lower lip, hairs
on them [ears] lacking light tips. Hairs on neck and trunk dark, red-
brown, guard hairs have broad (about 2 cm) light tips. Tail dark
brown, without light highlights. Limbs dark brown, without admix-
ture of light hairs, quite sharply delineated from the lighter trunk.
Claws are dark horn or blackish-brown color.

Dark type: general tone blackish-brown, sometimes with weak
light highlights (gray hair). Base of guard hairs brownish-black,
underwool brown. Muzzle dark, chestnut brown. Forehead and side
of head gradually lightens posteriorly. Ears dark brown without light
tips on guard hairs. Trunk and neck have guard hairs with very
short light tips. Belly and limbs are very dark, brownish-black.
Claws as in light type (Flerov, 1935).

Both types are connected with each other by a full series of
transitions, the dark type predominating (“brown” according to
furriers), especially in the western part of the range (in the middle
districts of the European section of the country and to its north,
bears are dark).

Greatest length of skull in males,?”? 311-М 325—388 mm, fe-
males—279—M 301—336 mm; condylobasal length of skull in males,

6Characteristics here and below employ data of Ognev, 1931; Flerov, 1935;
and Kuznetsov, 1952, with some changes and additions.
27Measurements here and below based on data of Ognev (1931).

430

637

300—M 320-343 mm, females—269—M 286-313 mm; zygomatic width
of skull in males, 117—M 207—231 mm, females—147—M 174—192
mm; length of upper tooth row in males and females—106—M 117—
125 mm.

Maximum weight 320—350 kg, as a rare exception, larger. Ani-
mals of the European part of the country are now smaller than
West Siberian, which is explained mainly by intensity of hunting in
the west. In the past century, this difference was less and very
large bears were encountered around Moscow.

European part of the Union, Urals, West Siberia approximately
to the Yenisei [river], and Altai (?).

Outside the USSR—Europe.

The eastern distributional border of this form is not established
and, apparently, is very indefinite. There is information that animals
of this race are distributed eastward, especially in northern Siberia,
farther than mentioned here.

2. East Siberian bear, Ursus (Ursus) arctos yeniseensis Ognev,
1924 (synonyms baicalensis, kolymensis; to this form the name
collaris is also given).

Dimensions larger than in Middle Russian bear (U. a. arctos),
but generally, considerably less than those in Far Eastern subspe-
cies. Particularly large individuals, however, often attain dimensions
characteristic of the latter. As with the latter, there are forms
corresponding to U. a. arctos or slightly exceeding it in size.** On
the whole, the range of individual variation in general dimensions
and skull measurements is very great—greater than that in U. a.
arctos and, apparently, greater than in Far Eastern forms.

Pelage long, dense and soft. Coloration generally of same type
as in Middle Russian form, U. a. arctos, but individuals of darker
color predominate (by terminology of furriers, “dark brown’).

Greatest length of skull of male is 326—431 mm; condylobasal
length of Ки 312-385 mm; zygomatic width is 193—252 mm; length
of upper toothrow is 121—143 mm.

East Siberia, beginning approximately at Yenisei [river], as far
as Trans-Baikaliya, Stanovoi range, Lena and Kolyma basins, and
generally throughout Yakutiya; Altai (?).

?6Namely these small bears were qualitatively described as special forms
(baicalensis, kolymensis). This notwithstanding that within the limits of the broad
range of these subspecies, there are small populations of the small bears.

638

Outside the USSR—northern part of Mongolian Republic. The
described form has been, so far, insufficiently studied entirely.

Ognev (1931) accepted three subspecies for the territory out-
lined above, giving them completely contrasting characteristics.
Adlerberg (1935) and Flerov (1935) assigned animals in the de-
scribed territory to U. a. arctos. The same was done by several
of our other authors, following them without special study of the
question (Bobrinskii, 1944; Novikov, 1957).

The actual situation, apparently, is consistent with the view that
the broadest part of Siberia is inhabited by a population occupying
an intermediate position (“transitional”) between European-Sibe-
rian U. a. arctos and the group of Far Eastern races. The picture
of character distribution in the sense of their maintenance in popu-
lations and, in part, their distribution within the territory, is
complicated. The normal type of bear here is larger than to the
west, and extraordinary large animals are encountered which are
not inferior, or only slightly inferior to typically large individuals of
Far Eastern bears. These animals, in part very old, occur, more-
over, not only in the east near the range of Far Eastern forms, but
also in the west—in the Yenisei basin (particularly in its upper
reaches), and along the Vilyui. Beside animals of this type, through-
out the whole region individuals are met with, not differing in skull
dimensions from U. a. arctos, or slightly differentiated from them.
They even appear far to the east (in Anadyr territory; Ognev,
1931), coming together here with typical Far Eastern animals
(piscator).

Altogether, this gives a picture of a transitional zone, where —
characters are related both with intraspecific hybridization, as well
as, apparently, a gradient [cline] between them. The particular com-
plexity of the picture is probably also caused by the presence of
transition between not two, but three, races: the nominal with the
two Far Eastern forms—the northern (piscator) and southern
(Jasiotus), which themselves introgress with one another (see
below).

Assigning animals of the described region to the nominal form
is, apparently, not any more, or only a little more, founded than
assigning them to one or another Far Eastern [forms]. Since the
range of bears of the above-mentioned type does not itself repre-
sent a narrow “transitional strip”, but is very broad (all of eastern
Siberia), there is a basis for recognizing them as separate subspe-

431

639

cies, perhaps of hybrid origin, and naming them. However, the
uncertainty of morphological characteristics of the race makes the
limits of the range distribution poorly defined, especially in the west.

The described form is analogous, in several respects, to the
Caucasian bear U. a. meridionalis (see below).

Note: In contemporary foreign literature (Ellerman and Morrison-
Scott, 1951; Haltennorth and Trenze, 1956) the form U. a. collaris
Cuvier et Geoffroy, 1824 is frequently mentioned as Siberian. Pocock
(1932) reestablished the forgotten and poorly defined name. The
range of this form is given as “Siberia”, in the upper Yenisei
(Ellerman and Morrison-Scott, 1956 did not give its overall range).
The forms yeniseensis and sibiricus are assigned it as synonyms.
One of the main characteristics of the subspecies is designated as
the well-developed light collar (Pocock, 1932). At the same time,
in the upper Yenisei and in Siberia, there are no populations which
could be characterized by a light collar, although there might be a
more or less significant percentage of such individuals. Everywhere,
this is a rare individual deviation. Moreover, the epithet collaris
applies to it. Therefore, this character does not offer the possibility
of establishing a type locality. Its indication as “Siberia” is too
general. In Siberia, as it is usually understood by foreigners, there
are three races in which a collar is occasionally found. In the
beginning of the nineteenth century, “Siberia” most probably re-
ferred to western Siberia, i.e. the region of occurrence of U. a.
arctos. The name collaris must, apparently, be considered a syn-
onym, or must not, in general, be given any nomenclatural significance
(nomen delendum).

3. Caucasian bear, Ursus (Ursus) arctos meridionalis Mid-
dendorff, 1851 (synonyms caucasicus, dinniki, smirnovi, ? persicus;
the designation syriacus was also applied to this form).

Dimensions moderate and small, very variable in size. In gen-
eral, considerably smaller than Middle Russian bear, U. a. arctos.
Pelage slightly fluffy. Coloration lighter than that in Middle Russian
form (U. a. arctos) and varies from straw-colored to light brown,
quite monotone—the range of its variation is less than U. a. arctos.

“General tone reddish-brown, comparatively light, without ad-
mixture of black and brown tones. Light frosting of tips of guard
hairs quite weakly expressed, and only on anterior part of body.
Underfur, light or light-chestnut. Guard hairs are brownish-reddish,
with very short light tips or without them. Limbs and abdomen

432

640

same tone as body and not darker than it. Muzzle somewhat lighter
than trunk, tips of ears same tone or slightly lighter than trunk.
Sometimes, collar occurs, developed slightly more strongly than in
U. a. arctos, in the sense of covering the neck. Coloration of some
individuals entirely monotone. Tail same tone as trunk” (Flerov,
1935). On withers, where hair is greatly elongated, a dark area
(spot) is evident, emphasized by somewhat darker color of under-
fur. Here, thanks to “disarray” of guard hairs, it [underfur] is more
easily seen.

Light and dark color types not sharply manifested. Darker
animals encountered relatively rarely, and deviations from more
broadly distributed, above-described type fall primarily on the side
of further lightening.

Greatest length of skull in male is 303—394 mm; condylobasal
length, 288—353 mm; length of upper toothrow, 110—133 mm.

Great Caucasus, Trans-Caucasus except in south and south-
west and Talysh—on the south including northern Armenia and
excluding coastal district of Black Sea northwards to Sukhumi
region.

Outside USSR, absent.

The Caucasian form of bear is very unique and of great inter-
est. Various, entirely different, opinions have been expressed about
it (Middendorf, 1851; Dinnik, 1914; Satunin, 1915; Smirnov, 1916;
Ognev, 1931; Adlerberg, 1935, Flerov, 1935). The extreme points
of view on one side assume the presence of four forms in the
Caucasus (Smirnov, 1916; Ognev, 1935), and on the other assigns.
bears of the entire Caucasus to one form; U. a. syriacus (Adlerberg,
1935).

While there is comparatively little color variation, the Caucasian
bear is distinguished by very great and complex variation in general
dimensions and skull; the latter are extraordinarily variable both in
size and structural characteristics. In the Caucasus, there is a very
large bear, not smaller, or only a little smaller than in the north,
having a large, massive skull with sharply developed protuberances,
crests—etc. of the same type as in the Middle Russian U. a. arctos.

Except for Pocock’s (1932) paper, no original studies deserving of attention
are found in foreign literature. Authors commonly agree with the viewpoint of one
or another of our investigators, often Adlerberg (1935), probably because it is the
simplest.

641

This is the type “caucasicus Sm.”. Its similarity to the northern
type is so great, that some authors (Dinnik, 1914; Satunin, 1915)
accepted the existence in the Caucasus of the nominal (U. a.
arctos) northern form of bear. Greatest length of skull of male of
this type is 348—М 368—394 mm. Some individual skulls are even
larger than in the eastern Siberian U. a. yeniseensis. Equally with
this, in some places in the Caucasus, there is a small animal, having
a small skull of the infantile type—with large braincase, weak crests,
narrow zygomatic arches, etc. (type “meridionalis Midd.”), living
side by side with them. Between these extreme forms, all transi-
tions occur (in part this is type “dinniki Sm.”). Such sharp
qualitative and quantitative variation gave grounds, with a purely
morphological approach, to their understanding of species and race,
for the description of various forms of bears in the Caucasus.

In the distribution of these types, there are some geographical
regularities. In the Main Caucasus range live bears of all three
types. They are found together (attempts to find ecological differ-
ences between them were unsuccessful). However, large animals
(caucasicus) occur more frequently in the middle, and especially in
the western, parts of the range. In the Trans-Caucasus, the large
form is absent, and bears met with here are of moderate and,
mainly, small (“meridionalis”) types.

Several geographic tendencies in color also exist. In the Main
range, together with light, a significant number of relatively dark
animals are encountered. In the Trans-Caucasus, such dark animals
are few, and light and very light predominate. In addition, there is
no sharp geographic localization, and mutual exclusion of different
geographic types. This is one race, which is, however, entirely
variable, not only in qualitative, but in part in quantitative, features.
There are only demonstrated tendencies in geographic displace-
ment of types of variation (see below, description of Syrian bear).

This picture of characters and their distribution in the population
directly indicates that the Caucasian bear, U. a. meridionalis, itself
represents a transitional, apparently, hybrid race from the Euro-
pean-Siberian group of bears (arctos), namely U. a. arctos, to the
Near Eastern group of bears (syriacus), namely U. a. syriacus.
Named thusly, this should, probably, decide the “enigma” of the
Caucasian bear.

Clearly unfounded is acceptance of several forms, as was al-
ready previously shown. However, so too unfounded is assignment

433

642

of all Caucasian bears to U. a. syriacus. As this form is not badly
known, it is certain that bears of the Great Caucasus as a popu-
lation have very little in common with it. They differ from it, and
from animals of the greater part of the Little Caucasus and Trans-
Caucasus (except the south; see below, description of Syrian bear).
As for the detail, that in the Trans-Caucasus there occur large
numbers of smaller and lighter animals, and few large and dark
ones, this is entirely regular. In the Trans-Caucasus, the features of
transition to U. a. syriacus are more evident, and the genetic
induction [sic] of this form is naturally greater than in the Great
Caucasus.

The data given about character traits of U. a. meridionalis and
geographic tendencies of their distribution in the population are
useful in understanding its other characteristics and their origin. As
it is known, the Caucasus, and particularly the Trans-Caucasus,
received its fauna, which is forest-associated to a considerable
degree, from the Near East and Asia Minor or from Europe through
Asia Minor. Probably the colonization of the Caucasus by bears
also came from the south. Thence, apparently, small, light bear
penetrated. This possibility exists at the present time. On the other
hand, in the past, even recently (Holocene), large and dark bear
also penetrated into the Caucasus from the пов. This penetra-
tion came from the northwest—from the Don to the Kuban’.

The mixing of these two lineages by unlimited hybridization
gave the existing variegated morphological and geographical picture
of the race. With this viewpoint, the presence of large, dark bears
(as well as small), mainly in the Great Caucasus and especially шо
the west, and the predominance of small, light ones in the Trans-
Caucasus is explained. It is noteworthy that in the Caucasus there
are bears whose skulis do not differ from U. a. arctos. Therefore,
U. a. meridionalis is a transitional form between the northern—
European-Siberian—and the southern—Near Eastern bears. Its
“polymorphism” is the result of its hybrid formation. This form
occupies a considerable and zoogeographically distinctive territory,
well characterized, and therefore deserves to be considered in the
nature of an independent subspecies.

3°A similar possibility was already assumed by Smirnov (1916). Adlerberg
(1935) is completely wrong in his affirmation that “in any case .... there is no
basis for arriving at the conclusion reached by Smirnov, that the bears of the
Caucasus speak to the influence on such a fauna of boreal character” (p. 100).

643

The Caucasian bear is, to a certain extent, analogous to the
Eastern-Siberian U. a. yeniseensis; however, the whole picture
here is apparently more sharply manifested; in particular, amplitude
of variation is greater. |

Note. Adlerberg (1931) suggested elimination of the name
meridionalis Middendorf on the basis that Middendorf understood
under this name, not only the Caucasian bear, but also bears of
Asia Minor and southern Europe, and that the skull of a small form
of bear served as type. This author suggested the name caucasicus
Smirnov, 1916 be employed for the described form. From a nomen-
clatural viewpoint this argument is entirely unconvincing. Middendorf
(1851) gave the characteristics and distribution, in which he first
mentioned Caucasus, and the name given by him is formally irre-
proachable. Adlerberg’s mistake was repeated by other authors
following him (Bobrinskui, 1944; Novikov, 1956).

Near Eastern bears—group syriacus

4. Syrian bear, Ursus (Ursus) arctos syriacus Hemprich et Eh-
renberg, 1828 (synonym Jasistanicus).

Dimensions small—not larger than small individuals of the Cau-
casian bear (U. a. meridionalis); on average, somewhat smaller.
Fur short, sparse and coarse. Color very light, predominantly straw-
colored, legs not darker than trunk. Hair on withers long, having
gray-brown base and short light tips, from which a relatively dark
area is formed here. In some individuals, dark stripe extends along
spine. Main color types are two: darker, sometimes approaching in
color the light form of Caucasian bear (U. a. meridionalis). Ani-
mals of light type, which are a majority, are significantly lighter.
They often have whitish fur, with characteristic development of
straw yellow or sometimes even rosy tones (very lightest animals
occurring are similar in color to polar bear; Adlerberg, 1935).3! Claws
brownish-horn with dark tips.

There are no data on craniological characteristics and dimen-
sions. Weight not more than 190 kg (Armenia; Dal’, 1954).

Black Sea coast of the Caucasus northward to Abkhaziya
(Sukhumi region), southern Trans-Caucasus south of Lake Sevan

3'“White bears” which sometimes appeared in the arena of the Roman circus

were, apparently, such very light animals of this form (Satunin, 1915).

434

644

(in particular, Sisiansk, Megrinsk, Kafansk and Gortisk regions of
Armenia; Dal’, 1954), Karabakhsk range, districts west and south-
west of Sevan (slopes of Mt. Aragats [Ararat]), Talysh and
Kopet-Dag.

Outside the USSR—Palestine (exterminated), Syria (partially
exterminated), Lebanon, Asia Minor, Iran.

О. а. syriacus is a very poorly studied form overall. This
particularly applies to animals living within the borders of the USSR.
Direct comparison of materials from there with “typical” syriacus
has not been done by anyone. At the same time, differences be-
tween bears in our above-mentioned regions and the other forms in
the Soviet Union are adequately determined. The characters of U.
a. syriacus from other parts of the range are also poorly known.
Bears are extinct in a series of places, are generally rare, and there
are few in museums. The main character of the race is, apparently,
the very light color and small general dimensions of body and skull.

Animals from the Pri-Caspian province of Iran and Kopet-Dag
(and, probably, from southern Trans-Caucasus and Talysh) are af-
filiated by several authors (Flerov, 1935) to the Caucasian U. а.
meridionalis. Judging at least by bears from Kopet-Dag (Ognev,
1931; V.G. Heptner), this is incorrect. It is possible, moreover, that
animals from northern Iran and Kopet-Dag are not fully “typical”
compared to those of southern Iran and Syria. The most recent
investigations associate all the bears of Iran to the described form
(Misonne, 1960). The limits of distribution of this form in the Trans-
Caucasus on the north are not sufficiently clear. Forest districts in.
the mountains of northern Armenia (north of Lake Sevan) and
northwest of there were, apparently, occupied by large Caucasian
bears (Dal’, 1954). It is also not known hc w deeply into the Caucasus
from the seashore this bear is distributed, and which form inhabits
the Trialetsk, Adzharo-Imeretinsk ranges and adjoining places, etc.

5. Tien Shan bear, U. (Ursus) arctos isabellinus Horsfield,
1826 (synonyms leuconyx, pamirensis; the species epithet prui-
nosus was applied to bears of this form living among us).

Size is quite small or moderate. Pelage soft, long, shaggy and
somewhat sparse. Color very light, varying from light-straw (whit-
ish) to light grayish-brown. Coloration monotone, extremities and
tail not darker than trunk; ears very light, whitish, always lighter
than general color tone. Developed collar occurs. Claws light.

435

645

Two main color types exist, connected by a complete series of
transitionals. “Light type: general tone reddish-straw or brownish-
white with translucent light sandy—sandy underfur. Muzzle light,
straw-whitish, hairs on forehead have dark base. On neck, shoul-
ders, nape, and band along anterior part of spine, the hairs have
brown, quite dark, base, but lighter tips than on other parts of body.
Ears very light, tone of muzzle lighter than general color tone. Belly
darker than upper trunk, hairs here do not have light tips,

Dark type: general tone light, rosy-brown, extraordinarily ho-
mogenous with a well manifested light frosting. Muzzle is somewhat
more reddish than trunk. Ears very light in color, straw-whitish,
lighter than whole body. Belly slightly brown, but becoming darker
dorsally as are extremities. Tail same tone as trunk. Underfur quite
dark, rosy-brown. Hairs have small light tips (Flerov, 1935).

Accurate data on the dimensions of this form within the bound-
aries of the USSR are actually absent. An old male from the
Chatkal range had a 140-cm body length, height at withers, 90 cm;
length of hind foot, 20 cm. Greatest length of its skull, 365 mm;
condylobasal length of skull, 320 cm; zygomatic width, 195 mm
(Sludskii, 1953). In general, skull length may reach 383.5 mm;
condylobasal length—356 mm, zygomatic width—233 mm (Ognev,
1931; one specimen), 1.е. as in Middle Russian И. a. arctos. In the
population, very small individuals are more frequent in the south,
analogous to the small form of the Caucasian bear U. a.
meridionalis, but they are rare to much more rare than in the
Caucasus. The type “И. pamirensis” is assigned to them. Great-
est length of skull, 279.5 mm; condylobasal length 261.3 mm;
zygomatic width, 172 mm (Ognev, 1931).

Mountains of eastern half of Middle Asia—Pamiro-Alai system
and Tien Shan eastward to the Ш river.

Outside the USSR—contiguous parts of Pamiro-Alai and Tien
Shan in China (limits of distribution eastward not known), western
Himalayas from Chitral in west to Nepal in east, Karakorum (7),
Waziristan, Afghanistan.

U. a. isabellinus is a well-defined form, close to U. a. syriacus.
Their characters overlap and the differences of both forms are
insufficiently clear. Sometimes they are united as one (Flerov, 1935)
which is, apparently, incorrect. Their ranges in western Afghani-
stan and eastern Iran are separated from one another. In the
northeast, U. a. isabellinus integrades with the European-Siberian

646

U. a. arctos. In the Dzhungarsk Alatau lives an intermediate popu-
lation, and animals with several such characters (light color) are
encountered in the southern Altai (Sludskii, 1935). U. a. isabellinus
is also very clearly distinguished from the Tibetan U. a. pruinosus
in craniological features, and its placement in this group (in “spe-
cies” pruinosus; Ognev, 1931) is ungrounded. The boundaries of
the ranges of these two forms in relation to each other are not
known. In Tibet itself lives only U. a. pruinosus.

The southern (Kashmir) representatives of the race are, possi-
bly, in general somewhat smaller than the northern (Tien Shan);
however, the difference is negligible and the bears of Tien Shan
should not be considered a separate form (Jeuconyx) (Pocock,
1932, 1941).

Far Eastern bears—group piscator

6. Kamchatka bear, Ursus (Ursus) arctos piscator Pucheran,
1855 (the epithet beringianus is often. used with reference to this
form; see below).

Measurements very large. Pelage long, dense and soft. Color
dark, predominantly dark brown; lighter color type rare. Limbs
always darker than trunk. Skull very large and massive, with broad
forehead region, which is quite steeply elevated above nasal bones.
Nasal bone relatively shortened, nasal opening broad; cheek bones
very widely disposed. Articular processes very wide; teeth rela- .
tively small. Claws dark.

Dark type. General tone of fur saturated, dark brown; in ex-
treme cases blackish-brown with dark violet-bluish tint cv. even
almost coal-black. Admixture of dark chestnut tone on head, in
region of shoulder-blades and along back. Extremities dark-brown
with dark violet tone.

Light type. General tone of fur brownish-reddish and even
golden-yellow, depending upon width of light tips of hairs on occi-
put, shoulder-blades and back. Sides of the head, withers and limbs
dark, cinnamon-brown. On the whole, bears of this type colored
quite variably. Lightening of color may be more intensive, and may
cover great part of trunk; limbs always dark.

Between both color types, there are all transitional forms.
Extreme forms of the light type are encountered in the form of rare

436

647

exception, extreme forms of the dark type—much more often.

Maximum body length reaches 240 cm, height at the withers,
135—140 mm, weight 650 kg and perhaps more.

Greatest length of skull of males 326—M 403—436; of females,
325—M 357-397 mm; condylobasal length of skull of males, 348—M
372—386 mm, of females, 303—M 354—370 mm; zygomatic width of
males, 212—M 258—277 mm, of females, 200-М 216-242 mm; length
of upper toothrow of males, 133-M 136—145 mm and of females,
119-M 126—133 mm.

Found in Anadyr territory, Kamchatka, Karaginsk islands, Kuril
islands, Okhotsk coast southwards to Stanovoi range and Shantar
islands inclusive. Extension of range deep into the continent is not
known.

Outside the USSR—Saint Lawrence Island and the Bering Sea.

The Kamchatka bear is a well-characterized race, related to a
number of the largest forms of the species as a whole. It differs
from bears of the group arctos very visibly, although as is known,
it is linked with U. a. arctos through the east Siberian form И. a.
yeniseensis. The limits of the range of both forms are unclear; in
some places, for example in Anadyr territory, both very large typi-
cal U. a. piscator and also bears of smaller dimensions very similar
to U. a. arctos, are encountered. The relationship of U. a. piscator -
to the form U. a. lasiotus, which systematically is closest to it, is
not yet sufficiently clarified. This applies both to characters and
also to distribution.

The Kamchatka bear is also very closely related to several
bears of Alaska and northwestern America (gyas-middendorffii
group). It is a single group of closely related races, characterized
by very large size (maximum for the species) and the above-men-
tioned peculiarities of the skull. Relationships of all Pacific Ocean
forms among themselves is, apparently, much closer than is now
believed. In particular, assertions of differences in measurements
(American are larger), are, apparently, exaggerated (see below).

Note. Instead of the name piscator Puch., employed for the
described form by Ognev (1931), in the last decades several Rus-
sian, followed by foreign, authors (Adlerberg, 1935 a; Bobrinskii,
1944; Novikov, 1956; Ellerman and Morrison-Scott, 1951, and sev-
eral others) began to use the name beringianus Midd., 1853. In
this case, they refer to the Sibirische Reise of Middendorf, where
in the second part of volume II, in figs. 1—6, is illustrated a skull of

437

648

“var. beringiana”. As already noted by Ognev (1931), it origi-
nated from Malyi Shantar Island. Ognev suggested considering this
specimen as the type of the form, and consequently, the type local-
ity is (some authors refer to B[ol’shoi]) Shantar Island; Ellerman
and Morrison-Scott, 1951). Ognev (1931) also showed that Shantar
bears belong to the Kamchatka form, and therefore it seemed that
denial of the name beringianus was not logical.

Authors using the name beringianus put out of sight that the
description in “Sibiris[c]he Reise is the second description, the first
having already been given in 1851 and in it for “var. beringiana
the following was stated: “distributed in the north, and especially in
northwestern America and in the countries of the whole eastern
coast of Siberia, including Manchuria and the Japanese Islands”.
Therefore, Middendorf, under the name var. beringiana, envisioned
all large bears of the Pacific Ocean districts of both continents.
However, it is indisputable that under contemporary nomenclatural
rules, the name beringianus belongs to one of the forms of north-
western America, apparently, in Alaska (it must, seemingly, replace
the name middendorffii).

The first name after 1851 may be considered that of Pucheran-
piscator—in 1855. This name is more suitable because it was given
to the population of Kamchatka, where the animals are very typi-
cal. The Shantar islands are located on the very edge of the
subspecies ranges, and the beginning of the region inhabited by the
form Jasiotus, where a mixture of the two forms are noted.

7. Ussuri bear, Ursus (Ursus) arctos lasiotus Gray, 1867
(synonyms mandchuricus, cavifrons; the species epithet yessoensis
was applied to some populations of this form).

In all respects similar to the Kamchatka bear, U. a. piscator,
but distinguished from it by somewhat more elongated skull, less
elevated forehead, somewhat longer nasal bones and relatively less
widely separated zygomatic arches. Teeth more massive, second
upper molar more elongated. Fur color as a whole somewhat
darker—light-brown to black.

Greatest length of skull of males, 387-М 423—455 mm;
condylobasal length of skull, 360-М 394—418 mm; zygomatic width
236—M 251-277 mm; length of upper toothrow, 137-М 145—151 mm.

Found in Ussuri territory, Sakhalin, Pri-Amur, northwards to
latitude of Shantar islands and, perhaps to Stanovoi mountain range,
and southern Kuril islands.

649

Outside the USSR—in northeastern China (former Manchuria),
Korean Peninsula, and Hokkaido Island.

Attribution of bears of the Mongolian Republic to this form
(Ellerman and Morrison-Scott, 1951) is incorrect.

Differentiation of Ussuri bears is not great, but real. Animals
may attain, apparently, a somewhat larger size than even Kamchatka
[bears].*? The separation of Sakhalin bear as a different form (Ursus
yessoensis, Ognev, 1931) is, probably, not justified. Range limits, in
respect to the Kamchatkan U. a. piscator and eastern Siberian U.
a. yeniseensis are not clear. Apparently, sufficiently wide regions
of transition exist in Pri-Amur, and perhaps, in Trans-Baikaliya.

It is sometimes asserted that in Ussuri territory and in Pri-
Amur where the above-described form of brown bear is found
together with the white-chested (black) bear (U. thibetanus), hy-
brids of these forms occur. This opinion is mistaken, and is based
on there being, among Ussuri brown bears, almost pure black indi-
viduals (Bromlei, 1956).

Outside the borders of the USSR, in the Old World, except for
those enumerated which are within our borders, and U. a. crowtheri
Schinz, 1844 (see above), there is only one other race—U. а.
pruinosus Blyth, 1854—of Tibet, western parts of Gansu and
Sichuan, Kam, eastern Himalayas, and extreme southern M[ongolian]
P[eople’s] R[epublic] (?).

Concerning American races of bears, they are discussed above
(V.H.).

Biology

Population. Within the boundaries of its contemporary range, popu-
lations of brown bears are distributed very unevenly. This depends
not only upon the extent of forests in separate parts of the range.
The bear is not simply a forest animal but an animal of large forest

The largest skull of the described form given by Ognev (1931) is only
slightly less than the maximum size of the world record (Alaska, according to
Haltenorth and Trense, 1956). If we take into consideration that in our country
up to now, “records” are not given any attention, and the hunters, and especially
traders, do not generally collect skulls as trophies, it is possible, perhaps, to
consider that animals of the described form (and also Kamchatka) attain dimensions
of large bears of America).

438

650

massifs. Its numbers depend also upon the degree of forest devel-
opment by humans (disturbance), and also upon differences in food
productivity (in general and by season). Of great significance is
direct pursuit on the part of humans.

Data on present populations are insufficient, and of an approxi-
mate character. Figures on the state of tanning are not indicative
of numbers, since brown bears are taken episodically, and their
skins often remain with hunters (no fewer than 60%).

In 24 districts in the middle zone of the European part of the
RSFSR, the minimal number of bears is about 4,000 (S.G. Priklonskii
and V.P. Teplov). The territory investigated covers, in the north,
Kirov and Kostromsk—in the south—Lipetsk and Tambovsk, in the
West—Smolensk, Bryansk and Kalininsk districts, and in the east—
Udmurtiya and Tatariya. (The above-mentioned figure is obtained
by extrapolation for the whole forested area of these districts from
a figure of 2,058 individuals; this latter was obtained by summing
data of a series of leskhoz [forest cooperatives], with an adjust-
ment of 25-30% for the possibility of duplication of the data of
mixed Jeskhoz). Ninety percent of the above-mentioned population
applies to the northern zone of this territory. Of these, 54% applies
to northern Kirov district and Udmurtiya, and to eastern Kostromsk
district. About 24%—to the Trans-Volgan part of Gor’kovsk,
Ivanovsk and Yaroslavsk districts, and to Chuvash and Marii ASSR.
The remaining 22% fall within the northwestern part of Smolensk
district, Kalininsk district, and Nekouzsk region of Yaroslavsk dis-
trict.> Very cautious extrapolation of typical densities gives 10—12
thousand for the districts of the European north, and for the districts
of the northwest, about 6,000 bears. In Estonia, about 180 bears
have been counted, and in Byelorussia, about 80 (Serzhanin, 1955).
Therefore, the estimated population of brown bears in the European
part of the USSR (excluding the Caucasus) hardly exceeds 22—23
thousand.

Population density (per 100 hectares of forest) is a very impor-
tant item in estimation of the bear’s living conditions. In the recent
past, the highest density was observed in Kamchatka, in the Kronotsk
ргезегуе—9—15 per 10 km?. Along a 10-km route in montane, berry-
bearing tundra, one may encounter up to 15 bears (Averin, 1948).

ЗА] these figures are very approximate and give only very general indication
of populations and. their sizes.

651

From 1940 to 1948, as a result of direct destruction, the bear
population in Kamchatka decreased catastrophically. A consider-
able density—from 1 to 3 bears рег 10 km? (1000 hectares) is
characteristic of montane forests of Sikhote-Alin’, the Caucasus
preserve on the northwestern slopes of the Main Caucasus range
(Nasimovich, 1940), the Pritelesk part of Altai preserve (Yurgenson,
1938), and also upland taiga on the right bank of the Yenisei [river]
(“Stolby” preserve). High density, with annual fluctuations from
0.4 to 0.9 per 10 km? is characteristic for the Central Forest pre-
serve (southwestern part of Kalininsk district; Yurgenson, 1937).

Moderate densities on the order of 0.25—0.40 are typical for
taiga forests on the upper Pechora and Ilych rivers (Teplov, 1960),
in the basin of the Konda and Sos’va rivers (data from Kondo-
Sos’vinsk preserve), southern Sakhalin (V.O. Shamykin), and for
the territory of the Altai preserve as a whole (G.D. Dul’keit). In
the European part of RSFSR, the forest along B[ol’shaya] Koksha
(upper Volga, Kalininsk district) represent an example of moderate
density—0.25 [per 10 km’] (Stroganov, 1934).

In 24 districts of the RSFSR, moderate densities have been
determined within a range of 0.03 (Meshchera) to 0.35—for the
northeast of the territory. For the northern zone of this territory, a
density of 0.20—0.35 is characteristic. For the Prisursk section, the
centre of Kirov district, and the southwestern part of the territory
(Bryansk district, southwestern part of Kaluga and Tula, and the
northwestern part of Orlovsk districts), typical density is on the
order of 0.10—0.17; for the central districts of the given territory—
0.03—0.08 (S.G. Priklonskii, V.P. Teplov). In Estonia, density reaches
0.15. An example of low density (0.12) might be the forest of the
Lapland preserve (О.Т. Semenov-Tyan-Shanskii).

In national parks of the USA (Yellowstone, Glacier) a bear
population density of 0.28—0.29 is estimated as moderate. In Roma-
nia, densities of 0.31 to 1.0 prevail; less than 0.3 is characteristic
for 1/3 of the hunting areas. Here there are hunting reservations
with a density above 1.1 (Almeshan, 1962).*

Habitat. Brown bear habitat within the boundaries of the USSR
is variable. In flatlands, it is, first of all, a forest animal; in the
extreme North, a periodic inhabitant of open tundras. In mountains,

*In Lit. Cit., given as 1963—Sci. Ed.

652

439. Fig. 155. “Bear crack” (Ayu-Ter)—habitat of Tien Shan brown bear,
Ursus arctos isabellinus Horsf. in the mountains of Tersk Alatau. Tien
Shan. Here, there are also Siberian ibex [Capra sibirica], gray marmot
[Marmota baibacina], and snow leopard (ounce) [Uncia uncia]. June

1964. Photograph by G.I. Zubanov.

it may be found both in forests, and also in high-montane meadows,
as well as among montane tundras. In Kamchatka, Sakhalin and
the Primor’e, a considerable quantity of bears restrict themselves
to the littoral of the sea coast at particular times of the year.

439

440

653

The bear prefers small-leaved coniferous forests. Among coni-
fers, it prefers forests with dark conifers—spruce, fir and nut pine,
but does not avoid pine woods and larch stands. This is explained
in that even in pinewood regions, dark conifer trees are closely
associated with taiga forest glades with numerous tall plants* and
also with tall-grass* vegetation along the banks of forest streams
and creeks, which serve as the main feeding places of this animal.
Owing to historical causes, the bear is linked to broad-leaved for-
ests only in the Caucasus, Primor’e territory and in parts of the
Carpathians.

Since activities of bears in their active period occupy an area
quite considerable in its extent, it cannot be restricted to any one
stand or type of forest, but only to a combination of them. This
occurs because each inhabited area must include feeding stands,
resting places, and migration routes. There are also winter den
sites, usually found outside the borders of the permanently inhabited
area.

Often, the various sites needed by bears are scattered in sepa-
rate sections among stable “migratory routes”. Thus, in the Pechora
taiga, in mossy, dark coniferous forests with berry bushes, traces
of bear activity amounted to 26% of occurrences, in pinewoods—
23%, in taiga glades with tall vegetation* along the banks of creeks
and rivers—28%, in open sphagnum bogs—8%, in birch groves—7%,
in grassy fir stands—4%, in unregenerated burns—2%, and in high-
montane areas—2% (Teplov, 1960). In the various months, the bear
prefers different biotopes: in April—May and in October—mossy, dark
coniferous forests and pinewoods, in June—July—August—taiga glades
with tall vegetation, then dark coniferous forests and pine groves;
and in September, again glades and sphagnum bogs.

Food. Geographic and seasonal variation is well demonstrated
in the food of the brown bear.

Spring is the most difficult and severe period occurring for the
bear, particularly early, until [growth of] vegetation begins. It is
particularly difficult for bears where they emerge from dens when
the snow cover is still deep (Kola Peninsula). In this case, the
animals in April and May feed mainly on ungulates (elk and

*In the Russian original the words krupnotrava and bol’shetrava both are used
here—what distinction the author wishes to make, if any, is unclear—Sci. Ed.

654

reindeer), attacking the living [animals] and eating carrion. Ants
and their larvae also occur as a permanent element in their food
(they excavate exposed ant-hills). In the upper Pechora, in April
and May, ants and their larvae comprise their main food (Teplov,
1960; P.B. Yurgenson). Remains of vertebrates and nut pine “nut-
lets” [seeds] are met with in small quantities in their excrement. In
the Denezhkin Kamen region in the Urals, cases were noted of
attacks on reindeer and hunting at capercaillie display-places.

In middle Sakhalin in spring, bears feed on the previous year’s
red billberry, ants, and marine flotsam (pinnipeds, fishes) and at the
end of the period—on shoots and rhizomes of tall grasses. On the
southern part of the island (V.A. Shamykin), marine flotsam is of
greatest significance (fishes, molluscs, holothurians), as well as in-
sects and maple twigs. In the Sikhote-Alin’ r[ange] in Primor’e in
spring, bears feed on the remains of the harvest of acorns, Man-
churian walnuts, and seeds of Korean nut pine (G.F. Bromlei, Yu.A.
Salmin and V.D. Shamykin). During famines, they feed on the red
billberries of the previous year. In years with poor harvests, besides
nuts and red billberry, they also capture insect larvae, wood-boring
ants and, later on, roots of saran (lily) and the first shoots of
graminoid vegetation. In Kamchatka (Averin, 1948), where the spring
period in the bear’s life lasts from April to about June, at first its
food consists of the previous year’s litter—especially stems of rough
blue-joint reed grass [Calamagrostis langsdorfi] and appearing
later, new shoots of willow, Silaus, meadowsweed [Spirea]
(shelomainika)* and large umbelliferous plants. The widely distrib-
uted reed-grass continues to be particularly significant, and, among
berries, the previous year’s cones and overwintering “nutlets”
[seeds] of prostrate nut-pine. As in Primor’e and Sakhalin, marine
flotsam, particularly the dead young ringed seals [Phoca hispida},
are of great importance in this period.

In the northeastern Altai, in April-May, the bears in forest glades
of sun-warmed slopes feed on roots of dog-tooth violet [Erythronium],
young shoots of puchok (cow parsnip) [Heracleum], angelica, cot-
ton-thistle [Onoporodon], and lungwort [Pulmonaria] as well as
horsetail and reed-grass (Yurgenson, 1938). In forests of the middle
zone, in April-May, serving as bear foods are the previous year’s
whortleberries (in mossy bogs), green shoots of young aspen and

*Local word?

44]

442

655

Fig. 156. Nut pines at the upper limit of the forest and tall-grass montane
meadows in mountains by Teletsk Lake—a feeding place for bears. Altai
preserve. Photograph by G.D. Dul’keit.

mountain ash, young sprouts of lungwort, and lichens from willow
bushes in mossy bogs of stunted trees—omshara* (Yurgenson, 1937).
In the northwestern Caucasus the main spring foods consist of
beech and oak mast (rarely chestnut and filbert), stems and leaves
of grasses, cow parsnip and coltsfoot, ants, beetles and their larvae
(Nasimovich, 1940). In some places, attacks on chamois occur.
A break in food comes on in May, when bear pastures appear.
The first period of summer, before the mass ripening of various
berries, 15 characterized by uniform feeding, in which stems and
leaves of tall forest vegetation are utilized—first of all umbellifers:
cow parsnip, garden angelica, wild angelica, and others. In the Far
East and Caucasus, coltsfoot [Pedasites] or bear burdock [Arctium,
Гарра] is eaten in great quantities, as is sow thistle [Sonchus] or
cotton-thistle, ramson [Allium ursinum]; and more rarely fireweed
[Epilobium], horse sorrel [Rumex confertus] (Pechora), cottongrass

*Local word—Sci. Ed.

443

656

[Eriophorum], marsh hawksbeard [Crepis paludosa], Sakhalin
calla, aspen leaves, and sprouts of huckleberry are widely utilized
and in great quantities. In the mountains of Sikhote-Alin’ bears strip
off the bark of the white-barked fir and feed on its cambium and
sap. As in spring, bears continue to breakup rotten stumps and logs
and eat wood-boring ants and their larvae as well as the larvae of
long-horned beetles, wood-boring beetles, and weevils. Geographi-
cal differences in feeding in this period are not great.

In the second half of summer, the essential role in the forest of
tall vegetation is retained, but fruit shrubs and bushes acquire great
importance. All over the whole taiga zone, and in forests of the
middle zone, huckleberry occupies chief place in bear’s food, and
in second place, raspberry, but the period during which it is eaten
is not prolonged. Among other berries, bears often eat blueberry,
rarely cloudberry (Pechora) and crowberry (Kola Peninsula,
Kamchatka), and along Sikhote-Alin’—berries of honeysuckle, yew,
Actinidia, Amur grapevine, and buckthorn; on southern Sakhalin
current [Ribes procumbens] and chokecherry. On middle Sakhalin,
salmonid fishes comprise up to 28% of the bear’s food in August.

In Kamchatka, from the beginning of July, among berries they
eat blueberry and crowberry (shiksha). Already beginning in July,
humpbacked salmon, salmon trout and other salmonid fishes ready
for spawning in the rivers by degrees acquire great significance.
Feeding on graminoid vegetation continues; some bears do not go
down to rivers at all and do not eat fish (Averin, 1958).

In the northwestern Caucasus, from the middle of July, rasp-
berry appears in the food of the bear, followed closely by blackberry
and current (Nasimovich, 1940). Later—green apples, cherry plum,
pear and sweetcherry. In rare cases, attacks on chamois were
observed. Significantly more frequently it feeds on carrion. In the
course of the summer, bears continue to feed on tall vegetation, and
eat bee:les, and ants and their larvae.

Autumn is the most important period in the nutrition of brown
bear since, particularly at this time, fat accumulation takes place
which is necessary for successful hibernation. In the majority of
taiga regions, it is secured by feeding on berries—whortleberry, red
billberry, cranberry, crowberry and others. In years of poor [berry]
crops in the Pechora taiga, late retreat into the den has been
observed (Teplov, 1953), wanderers are encountered and cases of
cattle attack become more frequent (60% of all cases).

441

657

Fig. 157. Sparse montane forest of nut-pines in Altai preserve. Pasturing
place of brown bears; in center of figure, under a nut pine, a bear is grazing.
Photograph by G.D. Dul’keit.

Typically, there is a transition from whortleberry and blueberry
to red billberry, cranberry and mountain ash in late autumn (from
October). The predominance of one species above the others is
determined by local conditions. Oats are also a typical early autumn
food. Its intensive utilization by bears in Kalinin district lasts for a
short time—15—20 days, from the first days of August when the oats
are still not fully ripe (Yurgenson, 1937); only rarely are bears in the
middle zone not attracted by oat. On the Pechora [river], however,
utilization of oats was not observed (Teplov, 1960). For fat accu-
mulation, the period of feeding on oat has, in some places, substantial
significance. In the middle zone, a lesser role is played in autumn
nutrition by acorns, hazel-nut, ants and graminoid vegetation. In the
Urals and in the Pechora basin, in productive years, nut-pine “nut-
lets” [seeds] gain importance. The same is found locally both in the

442

658

Fig. 158. Runways of water voles [Arvicola terrestris] uncovered by bear
in subalptne meadow. Tele massif, Altai preserve. Photograph by G.D.
Dul’keit.

flatlands and in the montane dark coniferous taiga of Siberia. With
insufficient food in late autumn, bears in the middle zone feed on
plant litter—aspen leaves, dry stems of reed-grass (Yuryenson, 1937).
In the oak-groves on the right bank of the Volga, acorns are of
great importance (V.A. Popov). In the alpine zone of the east[ern]
Altai, bears in autumn energetically dig out the long massive roots
of sweet vetch [Hedysarum].

Autumn foods of bears in the Far East have their own pecu-
liarities. In Kamchatka, in October-November, they eat “nutlets”
of prostrate nut-pine and mountain ash, and salmonid fishes—East
Siberian char and silver salmon. In years of famine, marine flotsam
gains particular importance, but berries and graminoid vegetation
remains of secondary importance (Averin, 1948). On middle Sakhalin,
bears in autumn feed on red billberry, “nutlets” of prostrate
nut-pine and migrating fish. On the southern part of the island

444

659

(V.D. Shamykin)—oats, migrating fish, marine flotsam, Actinidia ber-
ries, large-fruited sweetbrier, elder-leaved raspberry, shrub mountain
ash, rhizomes of coltsfoot and Urukundo maple shoots. In the moun-
tains of Sikhote-Alin’ (С.Е. Bromlei, Yu.A. Salmin and V.D.
Shamykin), from the middle of August, bears en masse eat various
berries, fruits of Maak’s bird cherry, Actinidia, Amur grapevine,
cork tree, buckthorn, yew, and “nutlets” of nut-pine. They eat great
amounts of spawning sima salmon [O. masu], humpback salmon and
chum salmon. In late autumn, the main foods for sustaining fat ac-
cumulation becomes nutlets of Korean nut-pine, acorns, red billberry,
in part blueberry from bogs and, of salmon—chum. Here, October is
the most favorable month for nutrition of the brown bear. In years
with deficiency of food, bears more often attack wild boar and roe
deer.

On the whole, during the active season of feeding in upper
Pechora brown bears, no less than 39 components of plant and
animal origin were revealed (P.B. Yurgenson). Of these, according
to frequency of occurrence (but not by volume), ants were in first
place—32.7%. Then followed fruits—21.3%, of which huckleberry—
12.7% and red billberry—5.9%. Vegetative parts of large-stemmed
plants constituted 34%, including umbellifers—10.4% (their rhizomes
only 3.6%). All mammals constituted only 9%, of which remains of
wild reindeers—2.7%; birds—2.7%, including capercaillie—1.3%. Of
other food, nut-pine “nutlets” constituted 5.9% of occurrences,
wasps—4.4%. Of the individual species of plants with edible green
parts; only one large hawksbeard [Crepis] (2.6%) and horse sorrel
[Rumex] (3.2%) gave occurrences of more than 2%.

For this period in the feeding of the Middle Russian brown bear
(Yurgenson, 1937), animal food constituted 65.4% of occurrences,
but according to volume only 38.4%. Invertebrates in this way gave
52.2% of occurrences, and vertebrates—only about 13.6%. Among
them, the most frequently occurring were ants—30.7% (by volume
only 9.1%). Among plant foods, raspberry occupied the first place
in occurrence—44%; then umbelliferous plants—27.7% (by volume—
52%). A little lower was the occurrence of oats—23% (by volume,
about 50%). Grasses gave 23% of occurrences, and aspen leaves
and shoots—9%. This is only a very approximate presentation of the
significance of individual food components. In Ussuri territory
(G.F. Bromlei), plant food comprised 68.9% of occurrences and
animal—31.1%. Among the former, berries comprised 33.9%,

660

graminoid plants—13.3%, nuts—10.1%; among animal food, insects—
16.8%, fish—10.1% and mammals—4% of occurrences.

In Romania, 28 species of plant food and 7 species of animal
food were shown for the brown bear (Almeshan, 1962).

Home range. Adherence to a defined home range is charac-
teristic of the majority of brown bears. Such ranges exist even in
those cases when bears continuously move widely from one place
to another in search of various seasonal foods during the course of
the whole period of activity. These movements are not disorderly
and are repeated by each animal from year to year in given areas.
These quite extensive vast and not strictly limited ranges are par-
ticularly typical for bears of mountainous regions—Caucasus, Altai,
Sayan, Sikhote-Alin’ (Yurgenson, 1938; Nasimovich, 1940; Yu.A.
Salmin and V.D. Shamykin).

Bear often stays in a den, outside its permanent home range,
since suitable places are not available everywhere. In winter, in
sections where there are many places for dens, the concentration
of bears may be very great.

Home ranges are well marked, where all conditions necessary
for the existence of bear in the forests are concentrated within a
limited area—300—800 hectares (Flerov, 1929; Yurgenson, 1931).
Dimensions of the range depend not so much on its abundance of
seasonal food, but more upon the area of “migratory stations”
included within its limits.

Burrows and shelters. During the active period, the brown
bear behaves as a typical denning animal. In the northern Urals and
Pri-Pechora taiga, its lairs are often found in montane forest mead-
ows and in river floodplains. They are distributed both in open
places among tall grass stands, and in the shade of lone standing
tree. Lairs also occur along the steep banks of taiga creeks—on the
slopes of low, wooded ridges,* among young pines and birches in
old burns, and by pine stumps in dry pinewoods. Sometimes, the
bear lies up in an ant-hill which it has dug up, and when aggravated
by blood-sucking insects, excavates a small hollow in moist ground
within a dense riparian thicket of wheat-grass [Agropyron] (P.B.
Yurgenson).

In the forests of the middle zone, preferred places for lairs are
dry elevated zone of “shore ramparts” almost on the very brink of

*Parma, local word in Urals—Sci. Ed.

445

661

the water along remote forest streams, the borders of tall-grass
openings, the edges of small forest bogs—mossy bogs and also
widely spaced trees and glades with grassy openings, dense thick-
ets of raspberry, young spruce and birch, usually with rotten logs
(Yurgenson, 1936, 1937). Bear dens are often restricted to areas
of tall grasses and their borders in the mountains of the northeast-
erm Altai (Yurgenson, 1938). In the mountains of the northwestern
Caucasus, dens often occur either under overhanging rocks, or on
snow fields, and rarely in open alpine meadows (Nasimovich, 1940).

Winter hibernacula in forests of the middle zone often occur
scattered along the edges of mossy bogs, especially on islands
among them, along the shores of forest lake and rivers, with dense
growth of mixed forests, among windfalls and near logged areas
(Mel’nitskii, 1915). At the same time, the bear does not avoid the
vicinity of traveled roads and settlements.

In the upper Pechora (Teplov, 1960), bears prefer to den in
pinewoods of the Pri-Pechora lowlands, where snow remains for a
less extended time than in the dark coniferous taiga of the foothills
and mountains. Of 16 hibernacula, 8 were constructed under over-
hanging slopes and logs, 6—aunder the roots of large nut-pines and
spruces, and 2—under the roots of young firs and pines.

In the mountains of the northeastern Altai and Sayan (Solov’ev,
1921; Yurgenson, 1936, 1938) hibernacula are predominantly “оп
the ground”, or located in natural caves, rock clefts, and in depres-
sions beneath rocks. Places suitable for this purpose occur rarely,
and therefore, groups of dens are frequently occupied by bears for
many years in a row. Thus, at Teletsk Lake (Azhi range), within
10 km, 26 dens were found. The majority of dens were restricted
to the crests of ranges, the upper parts of slopes, and inaccessible
ravines of montane creeks.

In Sikhote-Alin’, a better place for a hibernaculum is within the
high-montane fir-forest on the northern slopes of hills and on the
summits of ranges (Yu.A. Salmin and V.D. Shamykin). In
Kamchatka, dens occur in very different places—among large rock
fields, under roots and overhanging birch, and in caves (Averin,
1948). Early retiring animals and females build dens more carefully,
with a special nest. In Siberia, dens are constructed in montane
ravines, in thickets in stands of nut-pines, in coastal precipices, and
in the roots of trees. The “bed” is made from the twigs of conif-
erous trees, leaves, grass, reeds and moss (Middendorf, 1851).

446

662

In the northerneastern Caucasus, the great preponderance of
bears den deep in the mountains in the dark coniferous forest zone
(Nasimovich, 1940). Here, dens most frequently occur distributed
along steep rocky slopes—in caves and rock clefts, rarely in holes
of large trees, and logs within rhododendron thickets. In the alpine
zone (rarely}—in deep caves among limestone rocks. As in the
Altai, in this region aggregations of hibernacula may be found—
“bear cities” “Above-ground” dens are constructed with one exit
in sandy ground, in rotten stumps, near dry stumps, pockets sur-
rounded by small trees, in wind-drifted deposits, under overhanging
rocks and in clefts in rocks. In making open dens, bears often
excavate tub-like pits with sheer walls which are covered by
branches with the tips pointing upwards; in this way, the height of
the den can attain 2 meters. Females with lonchak [yearlings]
(young born in the preceding year) prefer to den in large piles
which form an arch, while the pregnant female prefers dense,
impassible brushy thickets (Mel ’nitskii, 1915).

Nearly every bear lines the hibernaculum with moss, coniferous
branches, dry grass, or hay—any material at hand. The quantity of
the bedding occurring is sometimes very great. The “brow” of the
den does not always face to the south or southwest, as is some-
times supposed. Sometimes, the den opening is plugged with branches,
especially in the “above ground” dens having lonchak.
“Kurzhak’—hoar frost above the hibernaculum resulting from the
respiration of the animal—is not always observed. The vicinity of
the hibernaculum is for the most part “cleared”*; the bear breaks
off tree branches which are used for lining the den.

Daily activity and behavior. Most often, in taiga forests and
forests of the middle zone, brown bears are active at twilight and
night, but on rainy days they usually roam the whole day (Yurgenson,
1937). In the alpine zone of northeastern Altai (Yurgenson, 1938),
and in montane tundra on Kamchatka (Averin, 1948), bears are
often observed even in the daylight hours, but mainly morning and
evening. In the unforested mountains of the northwestern Caucasus
in summer, 0.22 bears per hour were observed from 8 a.m. to 10
a.m., from 10 a.m. to 2 p.m—40.14, and from 2 p.m. to 6 p.m—0.20
bears; only 0.11 adult bears were observed in the morning hours;
during the day 0.04, and in the afternoon—0.20 respectively

*Literally, “broken’—Sci. Ed.

663

(Nasimovich, 1940). Females with young.come out earlier in the
hours after midday. On cloudy, rainy days females often graze the
whole day. With the passing of autumn in the broad-leaved forest
zone, nearer populated points bears become nocturnal animals, hid-
ing during the day in thickets of azalea, rhododendron, laurelcherry
[Laurocerasus| and ferns or even retreat to day-beds 6-8 km
higher in the mountains. In the northeastern Altai (Altaisk preserve)
in summer time, bears to a great extent feed in alpine meadows
most often during the day. However, during periods of prolonged,
extended rain, which are common in these mountains, they do not
enter the highlands for many days, keeping themselves lower—in
the nut-pines. With the arrival of clear, warm weather, they at once
move in significant numbers to the meadows, where they feed for
the entire day as if compensating for the omission (V.G. Heptner).

Thus, the brown bear. lacks a strict rhythm of daily activity.

Despite its seeming clumsiness, the bear is silent, light and

quick in its movement. Seriously frightened, for instance after an
unsuccessful shot, it rushes at great speed with huge leaps, extend-
ing the hind limbs far forward, and very quickly turning to escape
(V.G. Heptner). In walking, its tracks are in a straight line, circling
only when fattening, and on approaching the hibernaculum prior to
entering into hibernation, but the animal always avoids obstructions
and cluttered places. Young bears willingly climb trees—in Primor’e
only until they are 1.5 years old (Bromlei, 1956). But where there
are many acorns, walnuts or wild bees, the bear does not lose the
_ability to climb its whole life. In the northeastern Altai, bears ob-
taining cones with nut-pine “nutlets” often climb to the tops of
nut-pines and break off their branches (Yurgenson, 1938). Such
damages have been noticed in other regions of Siberia. In Poland,
earlier, bears were even caught by snares set in trees; with the
decline of wild-bee keeping, bears ceased climbing (Middendorf,
1851).

Bear digs well—in autumn, in digging out holes and the under-
ground storerooms of chipmunks [Тат!а5 sibiricus], it excavates
deep holes, often in rocky ground. In alpine meadows of the north-
eastern Altai, it digs up long, thick roots of sweet vetch [Hedy-
sarum]. In the middle [forest] zone, in obtaining food, it sometimes
lifts up the turf and rolls it into a tube in forest meadows. In the
high-montane meadows of the Altai (Altaisk preserve), bears in this
way eat subterranean parts of plants and catch voles, “rolling up”

447

664

entire large “carpets” (V.G. Heptner). Bears swim well, over-
coming distances of up to 6 km and they eagerly bath in hot periods,
especially cubs (Yurgenson, 1936, 1937, 1938; Nasimovich, 1940).
In order to get rid of blood-sucking insects, they make mud-baths.
During periods of drought, they excavate deep holes in mossy bogs
in search of water.

As in adult bears, so also cubs rub against tree trunks, leaving
their fur as well as claw marks. As early as 1874, the suggestion
was put forward that the bear marks its home range by its claws,
giving notice to other bears that they were approaching it (Egornov,
1874); quite a bit later, this was also stated by E.T. Seton-Thomp-
son (1911). Even later, the suggestion was advanced that these
marks are connected with the breeding period, but this is contra-
dicted—fresh marks are found at various times; and females also
make them.

Usually, the bear attacks humans only when it is injured, guard-
ing a fresh kill or when extremely emaciated and hungry [a condition
called] shatun (in winter). A female, surprised with its cubs by a
man, most frequently tries to frighten him by rearing up [on hind
legs]. Sometimes, she jumps directly at the person. As a rule, the
bear, upon sees a human, flees. Exceptions exist in some districts
where they are not pursued by hunters.

Hibernation and winter sleep. A characteristic feature of the
biology of bear is winter sleep in a hibernaculum. This is not hiber-
nation as in marmots and similar species, but is a particular sort
of shallow winter torpor. In this period, the development of em-
bryos, birth and lactation proceeds in the female. The intensiveness
of winter torpor develops gradually, and in the second half of win-
ter, bears are “fast asleep”. A necessary precondition for entry
into sleep is accumulation of fat deposits in a quantity sufficient for
maintaining life in the inactive period, and also in the period of time
after emergence from the den early in the spring. In the hibernating
bear, the rate of breathing and heartbeat decreases somewhat, and
its temperature falls to 29-37°C. Oxygen demand and carbon
dioxide output are sharply decreased. The intervals between inha-
lations become more prolonged and less regular. After 5—10
inhalations and exhalations, a pause of 2 to 4 minutes sets in
(Lobachev, 1951).

Bears accumulating fat in timely fashion hibernate earlier.
Animals that do not have sufficient amount of fat do not hibernate

665

Fig. 159. First snow in Altai preserve on balds in Chul’cha basin. Place
traversed by bears while going to hibernation places. In foreground, track
of animal which has just passed. Photograph by G.D. Dul’keit.

at all (shatun) [see prev. pg.] grow extraordinarily thin, attempt to
attack animals and sometimes humans, and often die from hunger
and freezing.

A necessary act before entering into the hibernaculum consists
of complete elimination from the alimentary tract of food remains.
In winter, in the rectum, a tight “plug” of wool, dry grasses and
other undigested food remains.

Entry into hibernation is preceded by selection and preparation
of the hibernaculum. Just before entry into it, the animal obliterates
its tracks, and often hibernates before deposition of snow, or at the
time of snowfall, which assists in hiding its tracks.

Hibernation does not take place simultaneously even in a single
region. Besides degree of nutrition, timing of hibernation is influ-
enced by the yield of principal foods and the particular weather in
that year. Entry into the hibernaculum proceeds from the end of

448

666

September (vicinity of Krasnoyarsk) to the first days of January
(Caucasus; Nasimovich, 1940). Timing of hibernation is obviously
influenced by geographic position. Thus, in Kamchatka, the prepon-
derance of bears hibernate at the end of October—females with
young, earlier (Averin, 1948). In middle Sakhalin, bears hibernate in
the middle of November before heavy snowfalls, in southern Sakhalin
(V.D. Shamykin)—at the end of November—beginning of Decem-
ber, in Sikhote-Alin’ (Yu.A. Salmin and V.D. Shamykin, G.F.
Bromlei)—from the first half of November to the second third of
December. In the upper Pechora (Teplov, 1960), based on many
years of data, timing of entry into the hibernaculum ranged from 3
October to 3 December—on average about 30th October
(Middendorf, 1851).

In the Trans-Caucasus some bears, especially in years with an
abundant yield of nuts, do not hibernate on the southern slopes.
Mass hibernation here takes place in the middle of December. In
the northeastern Altai (G.D. Dul’keit), the period of entry extends
from the end of November to the beginning of December (400—
1000 m.n.u.m.)* and in the years with little food—from 9—10 to 20
December.

The duration of stay of the brown bear in the hibernaculum
ranges from 75 to 195 days per year; in the Caucasus—75—120, in
southern Sakhalin—135—145, in the eastern Sayan—145—165, in the
Kola peninsula and in Kondo-Sos’ vinsk ргезегуе—180—195 days. In
individual years, it fluctuates significantly. Thus, in the upper Pechora
(Teplov, 1960) it equals on average 165 days, with a range from 138
to 190. Therefore, geographic variation in duration of the hibernation
period differs somewhat from the multi-year average for a single
region.

Timing of emergence from the hibernaculum is determined by
various factors: well-nourished animals emerge later than emaci-
ated ones. Early emergence is also associated with an unsuitable
choice of location for the den (premature thawing), or disturbance
caused to the animal by hunters or tigers (Sikhote-Alin’). In the
Caucasus, on the southern slopes of the Main range, the bears are
aroused earlier than on the north; in Kamchatka in the interior of
the peninsula, later than near the seashore. Females with cubs are
the latest to emerge.

*metr nad urovenem morya; meters above sea level—Sci. Ed.

ecm. ad .

Plate 4. Middle Russian brown bear, Ursus (Ursus) arctos arctos L. Painting
by A.N. Komarov.

449

667

Bears in the Caucasus are the earliest to arouse, where the
earliest date recorded was 5 March, and the average date—14—25
March. The period of concurrent emergence occurs within 10—12
days, and females with cubs leave 9-10 days later (Nasimovich,
1940). Bears on the Kola peninsula are latest of all to leave—about
| May (О.Г. Semenov-Tyan-Shanski1). In Anadyr territory, bears
leave at the end of April (Sokol’nikov, 1927), on the Kolyma (Buturlin,
1913) and in Kamchatka (Averin, 1948) where the earliest time
was recorded on the 4 April and the latest date—the middle of May
(lactating females).

In different regions, there exist significant fluctuations of tim-
ing (Table 46). Thus, in the Caucasus preserve, the earliest date
observed over 10 years ranged from 5 to 27 March (Nasimovich,
1940) and, in one year (1950), the dates stretched out from 14
March to 18 April (S.I. Chernyavskaya). In Pechora-Ilychsk pre-
serve during 13 years (Teplov, 1960), date of first emergence ranged
from 27 March to 25 April (on average about 16 April). In the
upper Pechora (P.B. Yurgenson), emergence from the hibernacu-
lum is connected with establishment of favorable daily air temperature
and the formation of openings in the ice [of the river]; (with it
occurs the arrival of white wagtail and emergence of chipmunk). In
this period, thick snow cover also lies over the Kola peninsula. The
latest date (for well-nourished animals and lactating females) coin-
cides with the beginning of vegetation growth (first arrival of cuckoos
manifests itself as one of the indicators).

Therefore, the boundaries of the USSR, dates of emergence
from the hibernaculum range between 5 March and the middle of
May, falling primarily in April. The duration of the mass emergence
ranges from 6 to 15 days.

Seasonal migrations and transgressions. As a rule, brown
bears are sedentary animals, keeping quite strictly to the limits of
their home ranges. However, various types of shifts are character-
istic for them. Thus, brief local concentration of animals in places
of seasonal abundance of one or other food has been observed. In
spring and late autumn, middle Russian bears are observed in moss
bogs where they are attracted to huckleberries in years of heavy
yield. In summer, they are concentrated in hazel thickets, especially
in cut-over areas. Early in autumn, a significant concentration, in-
cluding the majority of individuals, is observed near the oat fields
adjoining the borders of forests; they are also noted in oak stands

668

in connection with the acorn crop (Gor’kovsk district, A.N.
Formozov).

More or less regular seasonal migrations are typical for mon-
tane regions. Thus, in the northeastern Altai (Yurgenson, 1938) in
the active season, bears ascend from the montane valleys to the
slopes three times and then descend again, in connection with various
dates of ripening of berries and nut-pine “nutlets”. In the north-
eastern Caucasus autumnal migrations are observed from interior
montane regions to the borders of the belt of broad-leaved forests
and their “fruiteries” (Nasimovich, 1940). These migrations are
characterized by great regularity and are associated with the fruit-
ing of oak, beech and other “mast trees”, and also fruits growing
wild in Circassian gardens. To hibernate, bears again move away
deep into the mountains. The distance of such migrations has been
determined as tens of kilometers. In the past, bears moved about
120—150 km (Dinnik, 1914).

Such wanderings appear in autumn in Sikhote-Alin’ (Yu.A.
Salmin and V.D. Shamykin; G.F. Bromlei) and depend upon the yield
of acorn, “nutlets” of Korean nut-pine and the distribution of yield

Table 46. Dates of emergence of bears from hibernacula

Region of observation Date
Lapland preserve About 1 May
Kolyma territory End of April
Anadyr territory Middle of April
Kamchatka 10 April—middle of May

Middle Sakhalin
Pechora-Ilychsk preserve
Denezhkin Kamen
(Middle Ural) preserve
Ural
Kondo-Sos’vinsk preserve
“Stolby” preserve
(Krasnoyarsk territory)
Northeastern Altai
(Teletsk Lake)
Southern Sakhalin
Aksu-Dzhabaglinsk preserve
Sikhote-Alin’ preserve

Leningrad and Novgorod districts
Central Forest preserve
Bashkir preserve

End of April
27 March—25 April
Middle of April

Second half of April
Beginning of April
8-28 April

Beginning of April—
beginning of May

Middle of April

End of March—Middle
of April

20 March—15 April

22 April2 May

3—11 April

450

669

in the area. With a failure of this main, but unreliable food supply,
bears here undertake significant migrations of up to 200—350 km.

In the upper Pechora and in the Urals (Sabaneev, 1878; Teplov,
1960) the movements of bears are associated locally with the low
yield of berries, and also, on the western to the eastern slopes of
the Ural range, is connected with later formation of snow cover
there. In Semigrad’a (Kaspaty; Yurgenson, 1936), massive regular
migrations from montane forests, repeated every 5 years, are pro-
voked by a heavy acorn crop in the oak forests of the plains.
Animals are drawn from the montane forests from a radius of 50—
100 km. Coincidence of abundant oak and beech mast crops makes
the migrations less intensive, but these are very rare. Bears do not
leave regions where the yield of beech mast and blackberry or red
bilberry is fruitful (Yurgenson, 1936).

Besides this type of migration, other migrations have been
observed which are caused by obligatory situations—major forest
fires and summer drought, causing the usual water sources to dry
up. Such cases were observed in the middle [forest] zone (Yurgenson,
1937) and in southern Sakhalin (V.D. Shamykin).

Reproduction. According to data of the Moscow zoo (Gulyaev,
1934), rut in the brown bears lasts about | month, beginning on 10
May. Individual cases of mating were observed in July. Copulation
lasts from 30—40 min. to 1 h. In case of absence of fertilization,
esterus 15 repeated after a fairly long period. Under natural condi-
tions, rut lasts from 10-20 May to the middle of July. In Chita
district (Pavlov, 1948), rut in bears coincides with flowering of the
coral lily (Lilium tenuifolium). The peak of rut at different geo-
graphic points takes place at various times within the mentioned
time limits but without the clear regularity which has sometimes
been mentioned (Middendorf, 1851). Among the signs of rut are
shown to be bellowing, fresh conflict, and bite marks on trees,
fighting (sometimes ending fatally for one of combatants), pursuit of
the female by several males, etc. In the upper Pechora (Teplov,
1960), these signs were not observed at all in June to July, but on
the contrary, occurred in the middle of September according to the
word of hunters. This coincides with data (Middendorf, 1851) ac-
cording to which rut extends for several weeks in July—August, but
also occurs in May and September. Duration of pregnancy is, there-
fore, 6 months. According to data of the Moscow Zoo (Gulyaey,
1934), it lasts from 185 to 251 days (8 cases); i.e. 6-8 months, 227
days on average (7.5 months).

670

Birth most often takes place in the first half of January (Gulyaev,
1934). Individual cases were observed in December, in the second
half of January, in February, March and even in April (Middendorf,
1851; Nasimovich, 1940; Stroganov, 1952). In the litter occur 1 to
5.cubs; more often—2, rarely 3; 4 to 5 cubs occur extremely rarely.
Young and old females often give birth to one cub (Middendorf,
1851; Gulyaev, 1934; Kuklin, 1937; Nasimovich, 1940 and others).
In 1958, in the Kirensk region of Irkutsk district, 5 cubs were found
in a den (Stroganov, 1962); in Valdai, 6 embryos were found in a
female (Shirinskii-Shikhmatov, 1900).

The sexually mature female bears a litter every other year,
since having juvenile cubs, they do not participate in rut. In Caucasus
preserve, about 35% of the females have offspring annually
(Nasimovich, 1940).

Growth, development and molt. The newborn cub weighs
501-510 g (Gulyaev, 1934), with a body length of 23—23.5 cm. At
the age of 14 days, weight has grown to 538—552 в, and body
length, 26—28 cm. Newborn cubs are sparsely covered with fur 2
to 3 mm in length, and their eyes are closed. The external auditory
meatus 15 covered by skin. The ear opening becomes defined on
the 14th day. Eyes open on the 30th—32nd day. By the third month,
it has a full set of milk teeth; by the fifth month the first molar
erupts, and in the sixth month, replacement of the milk teeth begins.
The last molar erupts at 10-12 months (Adol’f, 1949).

Lactation period lasts about 0.5 year. The cubs separate from
the female in their third year of life, having spent the first two
winters together with the female in a single den.

Bears grow slowly and full development and growth is not
attained earlier than 10 years, and according to some data, 20. A
series of changes in skull structure take place until very old age
(Zhitkov, 1922). Maximum age of bears in captivity is 47 years. In
Bern, a female gave birth to young in her 31st year of life.

Spring molt in the brown bear takes place slowly and 15$ рго-
longed (Kuznetsov, 1941). In the northeastern Altai, at the end of
April-beginning of May, bears are still found in winter fur. In June,
in the time of the breeding period, rapid molt takes place. However,
autumn molt proceeds slowly and unnoticeably.

Enemies, diseases, parasites, mortality and competitors.
Except for hunters, bears have almost no enemies. Primor’e terri-
tory constitutes an exception, where the Ussuri tiger may be an

451

452

671

enemy, most often attacking bears in winter, in the hibernaculum.
Since tigers are almost extinct, such cases are rare and do not have
any actual significance. In Caucasus preserve, in June 1931, it was
recorded that a leopard killed two small bears. However, this ani-
mal is also extremely rare here.

In Primor’e territory (G.F. Bromlei), lynx and wolf may be
considered enemies of cubs. There is information on attack by
wolves on bears in Kamchatka, but it is not reliable (Averin, 1948).

Competitors of bears are animals that are chiefly herbivorous.
Among them are included all consumers of forest berries and nuts,
ungulates grazing in forest openings with tall grasses, etc. However,
it is doubtful that the activity of these competitors might have vital
negative significance for bears.

Concerning the diseases of bears, there is no information. Sig-
nificant infection with endoparasites has not been recorded.

In an analysis of age composition of 58 bears killed in winter
in hibernacula on the upper Pechora (Teplov, 1960), newborn cubs
constituted 31% of the whole population, /onchak [yearling] (born
in preceding уеаг)—12%, and “pestun” (to 3 years)—5%. These
figures reflect, to some degree, the dynamics of bear mortality at
early ages. In the first year of life, it is equal to 61.2%, and in the
second—57.2%. On the whole, for the first years it is 85%. These
data cannot be presumed to be accurate, but they show that mor-
tality among cubs is very significant and explain the slow rate of
growth of the population. If, for the Pechora taiga, juvenile bears
were 31%, then in the Central Forest preserve (P.B. Yurgenson),
it was in different years from 17.3 to 33.3%, averaging 26.6%, and
in the northwestern Caucasus (Nasimovich, 1940), it corresponded
to 13.0-31.0%, 23.0% on average.

At an older age, mortality among bears, where they are not
hunted by man, is very small, and as a consequence, may attain
very advanced age and very large size. In Kamchatka where, in
the past, the largest bears weighed up to 640 kg (Averin, 1948), at
the present time, animals heavier than 300 kg are not found be-
cause of intensive hunting of them.

In the Central Forest preserve region, about 11% of the total
population are shot by hunters annually (P.B. Yurgenson).

Accidents are another cause of death among adult bears. Cases
are known when bears, falling into an opening in the ice, drown or
disappear under the ice of rivers and lakes. A series of cases are

451

672

Fig. 160. Tracks of northern brown bear, Ursus arctos arctos L., while

walking slowly. Tracks of hind feet do not conceal imprints of fore-feet.

Ponazyrevo, Kostromsk district. 5 November 1952. Sketch by
A.N. Formozov.

453

454

673

also known when male bears die during fights between one an-
other, and a case of the death of a bear during an attack on an
adult elk.

Population dynamics. Changes in number are characteristic
for brown bears, but they are poorly known. In the upper Pechora
(Teplov, 1960), the number of bears for 9 years fluctuated in a
relative index of 1 to 4; with 2-3 short fluctuations between con-
secutive years. These changes are explained by migrations in years
of berry failure. This reason, as well as forest fires and summer
drought, cause bears to wander, emigrating and changing numbers
in the forests of the southwestern part of Kalinin district (Yurgenson,
1937). Change in numbers 15 also caused by fluctuations in annual
population increase, together with changes in the quantity hunted
(including harvest). Thus, in the Central Forest preserve (Kalinin
district), over 19 years, a twofold change in numbers of bears was
observed. There was also a twofold fluctuation in litter size, the
number of females with litters, and the percentage of young in the
population also changed within these limits. When annual mortality
of bears due to hunting outside the limits of the preserve equaled
11%, yearly growth exceeded it more than 2 times. This, as well as
the small number of large animals, showed that a part of the popu-
lation dispersed every year. At the same time, new individuals
arrived which were previously not present. Locally populations in
the mountains of the northeastern Caucasus (Nasimovich, 1940)
have a nearly twofold fluctuation in the percentage of young
(13-31%).

Field characteristics. The brown bear is very vigilant and
cautious, and therefore is rarely seen by eye in forests on the
plains. However, its presence may be known by many signs of its
activity. This is first of all its feces and tracks, which are found on
the surface of damp or wet soil, on wet moss, and on snow. Bear
feces are extremely variable in color and consistency, depending
upon the food eaten. In size, they may be compared only with feces
of large domestic animals, but they are easily distinguished by the
incomplete digestion of the plant food.

Prints of fore and hind paws are sharply differentiated: forefeet
characteristically have prints with long strong claws, and also the
width of the track is equal to or greater than its length. Greatest
width of the track ranges from 9 to 19 cm. Tracks of the hind paws
resemble tracks of bare feet of humans (claws are not always

674

452 Fig. 161. Track of right hind foot of same animal in F ig. 160, on shallow wet
snow. Sketch by A.N. Formozov, about 2/3 natural size.

453

675

Fig. 162. Tracks of left fore and left hind feet of Tien Shan bear, Ursus

arctos isabellinus Horsf., on wet clay. Aksu-Dzhabagly* preserve in Talassk

Ala-Tau, 6 May, 1955. Sketch by A.N. Formozov after schematic of F.D.
Shaposhnikov, about 1/2 natural size.

noticeable), but are sharply differentiated by the narrow heel and
flat sole. Length of track of the hind paw ranges from 16 to 32 cm
and width, from 8 to 14 cm.

Presence of the bear in the forest is also betrayed by rotting
stumps and logs broken in its search for wood-boring ants; de-
stroyed anthills and nests of terrestrial wasps (not to be confused
with narrower and shallower holes of badgers); turf in forest open-
ings rolled in a tube; excavated burrows of chipmunks; severed
stems of umbelliferous plants (place of bite is ragged); trails pro-
ceeding through riparian or coastal tall vegetation beds in forest

*Misspelled “Dzhebogly” in Russian original—Sci. Ed.

676

openings and montane meadows; in autumn and spring, broken tips
of young mountain ash and aspen trees: ragged bites on young
trees; bark on trees stripped off by claws and deep claw tracks on
stems (sometimes with traces of wool}—‘“bear marks” (P.Yu.).

Practical Significance

The commercial value of the brown bear is not great; in the 30’s,
3—4 thousand skins were prepared annually. This figure does not
reflect the actual quantity of the catch of the animals, since a
considerable number of skins are retained by hunters for their
personal requirements. In 24 districts of RSFSR (European part),
out of 700 bears caught, the percentage kept was 60% (V.P. Teplov
and S.G. Priklonski1).

The warm, but heavy, skin of the brown bear, with its coarse,
thick dermis, is used only for preparing carpets, sleigh robes and
rarely for travel coats. Bear fat 1s utilized by the food industry and
is used for technical purposes. Its meat is also used as food, but not
everywhere.

Representations of the damage brought about by brown bears
to domestic livestock are greatly exaggerated. In a series of re-
gions, where bears are especially numerous due to favorable
conditions for their existence, it 1s just there the least harmful
(Kamchatka, Altai, Caucasus); it feeds mainly on plant foods and
almost never attacks livestock. But in those regions where loss of
livestock to attacks by brown bears is observable, it is caused by
only a few bears which develop, for one or another reason, the
habit of attacking livestock. The majority of animals are completely
harmless. Destruction of 1-2 of these predators completely stops
similar cases, despite the large number of bears here. On the con-
trary, an unsystematic hunt of the first animal encountered and shot
does not carry any kind of value.

Cases of attack by bears on humans are extremely rare. These
cases are often observed when the person turned out to be be-
tween a female bear and her cubs, or an encounter with a wounded
animal. Seriously emaciated, hungry “shatun” also attack humans;
1.е. bears which do enter into hibernation in winter, not having
accumulated sufficient quantities of fat, and sometimes also hungry
bears (especially early in spring) disturbed at a fresh kill (Yurgenson,
1933, 1936, 1937, 1938; Teplov, 1953).

455

677

Needless to say, ш a series of districts, hunting regulations
wrongly place brown bears among the number of harmful animals
which may be killed throughout the year. Carrion-bears most often
attack livestock only, and especially those found in fenced pastures
without herdsmen. In the taiga zone and locally in the middle [for-
est] zone, brown bears, in the course of a short period in milk stage
of oat ripening, feed on them, trampling down more than they eat.
This is not observed everywhere. Such cases were more frequent
in the past, when individual peasants, farmers and forest-guards
were sowing oats in small openings in the forest. There are a series
of simple means for frightening bears from oats—hanging old clothes
and rags in a circle around the field, stretching a string soaked in
kerosene, going barefoot around in the field at night in the dew, and
similar means. The oats bear is no more harmful to oats than
domestic livestock allowed to escape by herdsmen. Sometimes,
brown bears destroy forest beehives; such cases have caused losses
in Byelorussia up to the present time (Serzhanin, 1955).

Brown bears are hunted with shotguns, lying in wait at night
near a bait, or at the border of an oat field, often in a stand
constructed in a tree, and also through accidental meetings. Traps
for bears are less effective and are rarely employed. In montane
regions, bears are spied out and ambushed during times they are
grazing in the forest openings and alpine meadows. Winter hunting
of bear is quite widely pursued—in the hibernaculum or by the
chase. For hunting in dens, /aika [dogs] are used. This is mainly
an amateur hunting sport. Sometimes, /aikas are used when hunt-
ing on biack* trails in order to track, overtake and bring to bay the
animal until the approach of the hunter. This method is not wide-
spread due to the small number of /aikas trained for this task.
However, this method is the most suitable for destroying carrion-
bears which attack livestock, since its use makes it possible to find
a particular animal, which is necessary.

At the present time, hunting regulations of brown bear differ in
the individual republics, and districts differ in an even greater patch-
work often in no way justifiable. They often permit bear hunting
throughout the course of the entire year, but this cannot be justified
because it does not solve the problem of protection of domestic

*Not snow-covered—Sci. Ed.

456

678

livestock. In Krasnodarsk territory, hunting of brown bear is regu-
lated by issuance of licenses. In Estonia, it is completely prohibited.

Taking into consideration the size of annual growth of the fully
mature part of the bear population, the size of the annual catch
must not exceed 10% of the total number. In 27 districts of the
RSFSR, it was equal to about 35% (S.G. Priklonskii and V.P.
Teplov) which must be considered excessive.

Brown bear may remain an object of tourist hunting, and of

planned sport-hunting through license (P.Yu.).

Subgenus of Polar Bears
Subgenus Thalarctos Gray, 1825
POLAR BEAR, OSHKUI
Ursus (Thalarctos) maritimus Phipps, 1774

1774. Ursus maritimus Phipps. Voyage towards North Pole, p.
185. Spitsbergen.

1776. Ursus marinus Pallas. Reise durch verschiedene Provinzen
d. Russ. Reiches, 3, р. 691. Laptev Sea.

1792. Ursus polaris Shaw. Museum Leverianum, 1, р. 7. Renam-
ing of marinus.

1908. Thalassarctos jenaensis Knottnerus-Meyer. Sitzungsber.
Ges. naturf. Freunde Berlin, р. 174. Iena Island, King Karl
Land, Spitsbergen.

1908. Thalassarctos spitzbergensis Knottnerus-Meyer. Ibidem,
p. 184. Seven Islands, Spitsbergen. (V.H.).

Diagnosis

Dimensions large. Color monotone, white. Bare areas on lower
surface of feet (callosities) very small and almost entire sole of foot
densely covered with hairs. Facial portion of skull relatively long,
but shorter than in brown bear. Last upper molar tooth small, nar-
row, not elongated, and only slightly larger than anterior; last lower
molar small and considerably smaller than preceding tooth (V.H.).

*Proposed here. See below “Note” in section “Geographic Variation”

457

679
Description

The polar bear is a massive animal, of heavy build on quite long,
thick legs. Feet very large and broad, forefeet especially. Hind part
of body massive and strong; stronger than fore part, which is rela-
tively weak and seems to be laterally compressed (chest is not
broad), withers lower than croup. Neck very long (relatively longer
than in all other bears), very mobile, head relatively small, pointed,
with narrow forehead and small rounded ears. They are smaller,
and not so wide, as in brown bear; greater part of them hidden in
fur.

Eyes small, dark; naked part of nose tip, as well as lips, black.
Lips less free and mobile than in brown bear. Tail very short
(relatively shorter than in brown bear) and completely hidden in fur.
Claws relatively small, smaller than in brown bear. Those on fore-
foot larger than on hind, and their length along the curve is 6.5—7.0
cm (digit ПТ); length of claws on hind feet 1$ 5.0-5.5 cm (digits [-
IV). They are sharp, weakly curved, and pure black in color.

Lower surface of fore and hind feet thickly covered with hairs,
except for very small bare areas (smaller than in all other bears).
On forefoot, besides bare digital pads, there is a narrow transverse
callosity, much narrower than in brown bear, in region of anterior
part of metacarpal claws, and small rounded callosity on outer
posterior part of foot. Position and form of naked areas essentially
the same as in brown bear. On hind foot of polar bear, callosities
are same as on forefoot, but outer posterior one is elongated or
pear-shaped. Form and size of callosities are subject to certain
individual variations. Moreover, in connection with shedding of fur
(molt?), callosities are evident to different degrees (this is also true
of animals in zoological gardens). Lower surface of feet between
callosities covered with long hairs, which is characteristic of the
described species. They entirely cover these callosities, and in this
way, when the animal walks on snow and ice, it steps not on the
bare callosities area, but on a thick hairy cushion.

The winter coat of adult animals is very dense and compact;
however, it is relatively short, although hairs on posterior part of
body are longer than on anterior. There are no long hairs on with-
ers, and hair length in the shoulder region is all of 5—6 cm. On back,
at croup, their length reaches 8—10 cm. Hair on lower part of sides
and on belly greatly elongated (13—15 cm), also on rear side of hind

680

Fig. 163. Polar bear, Ursus (Thalarctos) maritimus Phipps. Sketch by A.N.
Komarov.

leg (13—14 cm), and especially on rear side of forelimbs (16—26 cm).
Very long, shaggy hairs (to 14.5 cm) outline the foot and protrude
in the region of the digits, covering them almost completely. On
bottom of foot, shorter, elastic hairs grow. Overall, fur on upper
part of body, neck and head short and lying compactly, and on
lower part of sides, belly and rear side of legs it is loose and
shaggy. Hairs, especially on upper body, quite coarse and elastic,
and on lower—flexible, depending mainly on their length. Overall,
fur is coarser than in brown bear.

Summer, 1.е. transitional, fur (one molt annually) shorter (on
belly about 8 cm, on rear side of forelimbs, 10—11 cm, on croup—
about 7 cm, etc.), but apparently, not sparse. Bear-cubs and young

459

681

animals have relatively longer fur coat than old ones. Geographic
variation in pelage is almost unexpressed.

Coloration is completely even on the whole body; white, some-
times pure white, often with slightly yellowish, sometimes strong
golden-yellow tint; in extreme cases, giving animal almost a sulfur-
yellow color.*> Sex variation in color is absent. Color of young is
usually purer white.

The skull of the polar bear is very similar to the skull of the
brown, but differs from it in the following characters: 1) upper
profile is straighter and projection of frontal region is not expressed
or weak; 2) facial region is somewhat shortened, and the length of
cranial region noticeably exceeds it: distance from anterior edge of
premaxillae to line connecting ends of supraorbital processes is less,
or, rarely, equal to distance from mentioned line to occipital crest,
and distance from end of nasals to mentioned line is approximately
two times less than distance from this line to end of occipital crest;
3) anterior part of muzzle is relatively wider: breadth of skull over
canines is greater than its breadth across infraorbital foramen (in
brown bear, such a ratio may only be found in large Far Eastern
bears—usually this ratio is reverse); 4) lower jaw is less massive,
and anterior edge of the last molar lies slightly behind level of
anterior border of orbit.

Canines are relatively strong, but molars are considerably smaller
than in brown bear. Last upper molar only slightly larger than first;
length of both molars is less than breadth of palate between first
molars (Fig. 150); width between outer sides of occipital and articu-
lar processes greatly exceeds length of upper molars and last
premolar of upper jaw (ratio is reverse in brown bear). Overall, by
comparison to brown [bear], dental system of polar bear has a
more “predatory structure”, in accordance with its mode of life and
nutrition. It is characteristic that even in very old polar bears, the
teeth are found in very good condition and are not at all affected
by wear. In brown bears, strong interaction of the teeth occurs and
wear begins very early; apparently, this is mainly related to the
plant food of the brown bear, and in particular with soil that occurs
on the teeth. Moreover, as may be judged from large series of
museum skulls, the appearance of acute caries and diseases of the

These yellow tones of the fur disappear during the dressing of hide, and bear
rugs always have a pure white color.

682

=
i
у,

TN ro
АХ RINT TG TT

=^ = Ss
Ee. 3
b ae” iN aS
aA As aay SI BY AN
YS See \\
BN IS У NV L
ЕЕ aN :
7 SSS SAS
> BSS №
ЕЕ:
Е #57
LY
233
ИЯ:
UALS LG; A:
SUI BA
ay ERS \ <) 7
— = \) YA
ВЕРА a ae
> у АВЕ

их
\\

G 3
Zi)

7222

OS
Z

(
Ss
=
77

=== == a UN
Ny
PNR

en 6 pe та №: SAN mS SN \ de
ou At, __

SM ee Loy Ui:

ay ty sj . ans

Lhe ; Wi My IW

р ssi ye ; БИ: sf КА
о Lins

Wars
ot

2) Ane
DOE Giro. SPURS

Ne A

siaietee’,

458 Fig. 164. Skull of polar bear, Ursus (Thalarctos) maritimus Phipps.

683

teeth, so frequently encountered in brown bears, sometimes with
serious diseases of the lower jaw, are almost not found in polar
bears.

Sexual variation is sharply manifested—the female skull is smaller
than the male skull, sagittal crests in them are shorter and weaker,
forehead is narrower, supraorbital processes consequently narrower,
and zygomatic arches are weaker. The skull of the polar bear cub
is similar to the skull of the brown, but differs by its narrower and
more protracted rostral part, and its zygomatic arches are weaker
and less developed.

Individual variation in the skull of the polar bear, both in dimen-
sions and in structural peculiarities, is relatively insignificant and
less by far than in brown [bear]. Overall, the polar bear is
craniologically monotypic.

Although individual, especially large, races of brown bears are
as large as the polar [bear], on the whole, the latter is larger than
the brown. Maximal dimensions of polar bears are larger than the
very largest brown bears, and they attain far greater weight.
Moreover, in its geographic variation there is no sharp contrast,
such as is found in brown (Kamchatka bear, U. arctos piscator,
Syrian bear U. arctos syriacus), and there are no populations in
which the types and range of individual variability are analogous to
those of the Caucasian, U. arctos meridionalis or the East Sibe-
rian U. arctos yeniseensis. Individual variation of the polar bear is
“normal”, but geographic variation is very small, which can be
explained by the basic conditions of its life and the type of its
geographical distribution area (see below).

Body length (without the tail) of adult males is 200—250 cm,
rarely to 285 cm, reaching 302 cm* as an exception; body length
of adult females is 160—250 cm; height at the shoulder is 130—140
cm, reaching 150 cm as an IAT ‘tail length (with terminal
hairs) is 20—22 cm.

Weight varies greatly, in part geographically, but chiefly in
accordance with season and degree of fatness. Weight of very
large males, in several parts of the range, is 700 kg and even 800 kg;

Information about a specimen 370 cm in length (Ivanov, 1933; after Tsalkin,
1936) does not merit confidence. This measurement often becomes exaggerated,
because authors usually measure the animals with the tail and terminal tail hairs
and sometimes the hide which is usually somewhat stretched.

460

684

there are data on bears weighing up to 1000 kg. Usually, weight is
less. Females are always not only smaller, but also considerably
lighter than males. Thus, males in eastern Greenland, in a status of
normal nutrition, weigh 400—450 kg, well-nourished to 500 kg; fe-
males in this region, even very well-fed ones, here rarely are larger
than 350—380 kg, 1.e. their weight comprises about 75% of that of
males. Particularly large bears have up to 112 and even 180 and
192 kg of fat (last two figures are probably exaggerated).

Greatest length of skull of adult male is 353—412 mm, females,
324—360.5 mm; condylobasal length of male skull is 359-388 mm,
females, 311—380 mm; zygomatic width of male is 187.5 to 274.5
mm, females, 194—199 mm; length of upper toothrow of males
110*—140 mm, females, 118—125 mm?’ (V.H.).

Systematic Position

The polar bear, in spite of its conspicuous external difference (color),
represents a species extremely close to the brown [bear], Ursus
arctos L. Only the American black bear stands closer to the brown
bear, apparently, than the polar bear.

One may consider, apparently, that the polar bear is, in a cer-
tain sense, derived from the brown, further increasing the
“predatory” trend (enlargement of canines, weakening of cheek
tooth row) in the series of species of the family. At the same time,
shortening of the facial portion of the skull, usual for the closely
[related] species, and which is considered as characteristic of a
certain sort of “primitiveness” (“infantilism”), here, apparently,
does not have this significance and may be interpreted as a special-
ized character. It is, apparently, connected with weakening and
shortening in the length of the toothrow. Moreover, the posterior
region of the skull has by no means any kind of infantile characters
(increased volume of braincase, basic weakness of crest develop-
ment, etc.), and externally is typical of large forms of brown bear.
In combination with other characters (coloration, dimensions, struc-

*In Russian original, “11’—Sci. Ed.

3’Measurements according to Hilzheimer 1930; Ognev, 1931; Birula, 1932;
Palkin, 1936; Heptner, 1936; Anthony, 1928; Pedersen, 1945, Shereshevski and
Petryaev, 1949. For additional data on measurements and their variation, see below
in section “Geographic Variation.”

462

685

ture of limbs etc.), in the polar bear is manifested a well-defined
complex of specialization for conditions in the Arctic, which is quite
obviously derived from the brown bear type. It is characteristic
that, having become an aquatic animal to a significant degree, in
any event swimming well and often, the polar bear does not have
the usual adaptations to the aquatic environment.

Palaeontological material, though not complete, also shows close
relationship of the polar bear to the brown, and allows us to. con-
sider it as derived from the brown. Thus, the dental system of the
polar bear of Eopleistocene time had greater similarity to that of the
brown bear than has the contemporary [polar] bear (last premolar
and second molar of upper [jaw]). In this way, the divergent evo-
lution of the polar bear noticeably advanced even in post-Pleistocene
time. Late Pleistocene U. m. tyrannus Kurt. was considerably
larger than the present form (Thenius and Hofer, 1960; Kurten,
1964) (V.H.).

Geographic Distribution

Shores, islands and floating ice of the Arctic Ocean in the Old and
New Worlds.

Geographic Range in the Soviet Union

The (reconstructed) range of the polar bear has a quite complicated
structure. In it four differentiated regions can be traced: 1) the
region of reproduction, by which is understood the territory where
females in winter lie in dens and give birth; 2) the region of summer
distribution of females and males of all ages; 3) region of winter
distribution (migration) of males and barren female bears; 4) the
region of long-distance movements to the south. With our store of
knowledge, it is impossible to outline precisely each region sepa-
rately.

The region of reproduction and the region of summer move-
ments, generally speaking, are bounded on the south by the southern
limits of the winter distribution region. In several places, the line of
the southern border of all types are quite close together, in others
they are quite distant from one another, often significantly. Here,
the southern boundary of the range is taken by the line limiting the
region of normal winter distribution of the animal. The region of
movements is considered separately.

"S89 OY} 0} UOISsoIssURy
Jo sjuiod эшецхо прапор oY} рие зэшц зиэтоце и! SoM OY} и! 991941950 JO 329 21415504—/ ‘291 и! paAlasqo элэм $1294 теоЧ
элэцА\ syulod Амэциои 330 `[елэлэ$—9 :(.орион,, ‘.ортехмон,, « зриеет [LINy,, ‘.UI[eyALS,,) 4910$ ay) 0} зиэшелош Jo ито
рэицэр AjIoog—¢ ‘adues [eUOU эцз JO зэмерипо4 oY} рио^эа YINos эцз 0} ззиэшелош JURISIP Jo Sade[d 2412945 — ‹(15рие]51) $1899
Зипох 0} Чиа эл! зэтещеу элэци ио18э1 Jo Alepunog—¢ :1894 JO эЗиел Jo Алерипо4 WOYINOS Чим эртэшоо JOU $э0р и элэЦи ‘5!
Зицео Jo Arepunog илэЧупоз jo эЗелэле Ieak-[NAI—Z ‘410$ oY} 0} зиоце18 ци [eWLIOU jo иотЗэ1 SUIpNjoul ‘эЗиел poyonsysuo0sal jo
Алерипод wayynogs—| :(owdeH ‘`О’А) USSN 9 и! “sddiyg snuuiapu (sojoapjoy]) упзаГ] ‘лез sejod ay) Jo asuey ‘с9т ‘9 197

051 оп
—————————

687

The southern (reconstructed) boundary of the range of the bear
in the Barents Sea is closely connected with the distribution of
floating ice. Its southern boundary includes the southwestern part
of the sea from the north, northeast and east. It begins at the
southwestern part of Spitsbergen, passes south around Medvezhii
[Bear] Island, is again southeastward approximately to 45° Е. Long.,
turns sharply to the southwest, reaching Svyat Nos in Murman
(multi-year average). In this way, the range included in early times,
Kanin, coastal tundras to the east of it (Timansk, Malozemel’sk,
Bol’shezemel’sk), the northern part of the White Sea and, appar-
ently, the easternmost part of the Murmansk coast.** However,
there are few data on this part of the range.

In the beginning and middle of the past century, the boundary
of the range, going on Medvezhii [Bear] Island, passed south of
Kolguev and Vaigach, but, apparently, did not include Kanin, and
probably, the coast of the mainland, except the Yugorsk Peninsula.
Thence, the southern boundary went eastward, tracking the coast
line right to the Bering Strait and to Provideniya Gulf. Beyond, the
boundary crossed to Alaska through the Bering Sea (see below).

The entire Baidaratsk Bay was undoubtedly occupied; in the
Ob’, it was, apparently, distributed north of the Arctic circle, 1.e. to
the mouth of Ob’; in the Yenisei Gulf, they were known up to
Gol’chikha and even to Tolstyi Nos in the very mouth of the Yenisei
at 70° N. Lat. North of this line, the range goes out to all the Arctic
islands and floating ice, to the Pole itself. The most northern point
of discovery is 86°04’ (Chapskii, 1946; there is information, not
very well defined, on the occurrence of bears at 88°15’).

If, in winter time, the southern boundary of the range shifts
maximally to the south (to the limits indicated above), then, as a
rule, in the far north bears are not met with north of 80’ N. Lat.,
occur there rarely, and do not penetrate far beyond this parallel. In
summer, on the contrary, the southern boundary shifts somewhat to
the north (very inconsiderably) but on the other hand, the range
strongly widens in the Polar region, and it is at this time that the
animals attain the Pole, i.e., they in fact live in the entire polar
basin. In that way, there is a seasonal, and in part very strong
displacement of the boundaries and of the whole range.

%8In exactly these regions polar bears were encountered by the people of
Veliki Novgorod* (‘“ushkuiniks”’).
*Now Novgorod—Sci. Ed.

463

688

The region of reproduction (reconstructed) is considerably
narrower than the general region of distribution and, within the
borders of our country, occupies only the Arctic islands—Novaya
Zemlya, Franz Josef Land, Severnaya Zemlya, New Siberian and
Medvezhii islands and Wrangel’ Island and small islands associated
with them. This part of the range includes also the numerous small
islands of the eastern and northwestern sectors and Kara Sea—in
the regions of Pyasinsk Gulf, Khariton Laptev Coast, Nordenskjéld
archipelago, Russkii and other more northern islands on the north-
ern Taimyr—Komsomol’skaya Pravda (Samuil) coast, Feddei, and
others. A great part of mainland coast of our Arctic area was not,
apparently included in the region of reproduction (birth of young) of
the bears—it includes only the extreme northern part of Taimyr,
north of latitude 76°, and on the east of the peninsula from 76° N.
Lat. to, probably, 74° N. Lat.*

At the present time, the boundary of the range has changed
significantly. There have been no bears in Kolguev and Vaigach for
a long time. In Novaya Zemlya, they are encountered along the
western coast southwards to the Admiralty Peninsula (middle of
the northern island), along the eastern coast, and also in the north-
ern half of Yuzhnyi [South] Island. Before the war, bears on Novaya
Zemlya were considered very rare. During the war years, their
number slightly increased, and the range, as related above, was
restored (S.M. Uspenskii).

Along the southern coast of the Kara Sea as far as Dikson and
somewhat farther to the east, the bear as a permanent inhabitant
is absent, and it occurs only as a transient in the very easternmost
part of the above-mentioned region. Already in the second half of
the 20’s at Dikson, and especially along the coast and islands of
Pyasinsk Gulf, and even more on the Khariton Laptev coast, it was
met with not only regularly in winter, but also in summer (V.G.
Heptner). It is frequently, and more or less normally encountered,
along the coast of northern Taimyr.

Along the coasts of the Laptev, East Siberian and Chukotsk
seas, the animals are completely absent in a series of sections, or
occur very rarely and accidentally. On the whole, it can be stated
that the southern boundary of the range is shifted northward and
bears at the present time are associated chiefly with the islands of
the Arctic Ocean—Franz Josef Land, Novaya Zemlya, Severnaya
Zemlya, New Siberian archipelago and Wrangel’ Island and with

*Not shown on map, Fig. 165—Sci. Ed.

689

the small islands associated with them. This took place, not only
due to the direct pursuit and killing of animals in connection with the
opening up of the Arctic in all its sectors (see “Population”),
beginning in the 20’s, but also due to warming of the Arctic during
the last 40 years, and particularly, 30 years. The shifting of the
boundary to the north is also observed in North America and in
Greenland, and climatic phenomena are also associated with this
(S.M. Uspenskii). Some increase in the number of animals, which
apparently took place in recent years in connection with its protec-
tion (see later) has not yet resulted in an increase (reestablishment)
in range. :

The region of reproduction, as regards its general outlines, at
least during the last century, has changed little and at the present
time, females den in almost all the area outlined above where it was
previously noted, except, apparently, several islands in the eastern
part of the Kara Sea. It is true that in Novaya Zemlya dens were
almost absent before the war (except perhaps the most northern
part on the Kara side); however, in the beginning of the 50’s,
females on Novaya Zemlya occupied dens, although they did not
range as far as in the past. In the 50’s, dens occurred along the
Kara side of not only Severnyi [North] but also Yuzhnyi [South]
islands (S.M. Uspenski1).

The main places of reproduction in our part of the Arctic are
Wrangel’ Island and Franz Josef Land. In each of these sectors,
150-200 female bears denned (S.M. Uspenskit, V.Ya.
Parovshchikov). In our remaining places, the number of dens of
pregnant female bears is less. For example, in Taimyr, there are,
in all, about 20 to 30. On the whole, over the entire Arctic there
are, apparently a total of up to 1000—1500 dens of pregnant females,
of which the majority (by several estimates up to 2/3) of which are
found in our above-mentioned region of reproduction (S.M.
Uspenskii).

Irregular and accidental transgressions of individual animals to
the south stretch out locally to very great distances beyond the
southern boundary of normal occupation. At the same time, bears
also appear deep into the mainland, far from the sea, but mainly on
the sea coast. In the majority of cases, these transgressions are
directly or indirectly connected with the transport of floating ice to
the south, with which the animals are very closely connected in the
summer time. They do not usually go beyond the limits of the

464

690

summer southern boundary of floating ice. Apparently, especially
distant transgressions occur in years of maximum ice, when the ice
is carried far beyond the limits of its multi-year average boundary.

Cases are known on European coasts of the appearance of
bear in Finmark, in Varanger fiord, and at Kanin Nos. These cases
are rare, occurring locally in the past [19th] and 18th centuries and
were, undoubtedly, connected with extreme ice years. In such years
of floating ice, the southern boundary of which lies in normal years
far from the shores of Finmark and western Murman (see above),
closely approach them. At Murman besides these [bears], appear
several cold-loving northern species which usually do not occur
here (Greenland seal, white whale; Heptner, 1930). Apparently,
under these conditions, the bears appeared. At the present time, in
connection with the extermination of the animal in the Barents Sea,
bears no longer occur along the coasts of the northern extremity of
Europe. Throughout the 20th century, it was apparently absent
here.

Cases have been recorded of transgressions into the mainland
on the Pechora from Ust’-Tsil’ma (about 320 km from the sea in
a straight line), along the Yenisei to Turukhansk (more than 500 km
from the sea on a straight line) and in Taimyr at the mouth and
lower reaches of the Khatanga, along the Kheta and in its divide
with the Dudypta (more than 300 km from the sea in a straight line;
V.H. Skalon), along the Kolyma as far as the mouth of the Omolon,
and others.

Along the Pacific coast, the animals were recorded in a series
of places in Anadyr Gulf, in particular, in its westernmost part at
the mouth of the Anadyr’, at Cape Navarin, along the coast of
Koryatsk Land (mouth of the Khatyrka, Shlyupochnaya Bay) at
Cape Olyutorsk, in Olyutorsk Gulf (Apuka) and on the eastern
shore of Kamchatka (Kronotsk peninsula, mouth of the Berezovaya
in the southern part of _Kronotsk Gulf). Farther to the south, wan-
derings are known to the Kuril islands, to the coast of Hokkaido,
and even to Honshu (Hondo; information not fully defined). In the
Okhotsk Sea, the polar bear is known to have been encountered on
the western coast of Kamchatka (Tolbachik village in Mil’kovsk
region), at Ol’skoe Island in Tauisk Gulf, near Okhotsk, on Sakhalin
(indeterminate reference) and even in the extreme northern part of
the Sea of Japan—at Mayacha anchorage near Sovetskaya Gavan’.

Accidental occurrence of animals on the Kuril islands and even

466

691

to the south in the Okhotsk Sea appears to be exceptionally rare,
and is completely irregular. The same applies, apparently, to the
eastern coast of Kamchatka. The appearance of bears in Koryatsk
Land and especially in Anadyr territory, though rare, may be con-
sidered an entirely regular phenomenon. It 15 certain that, in previous
times, at the time of large absolute numbers of animals in the north,
it appeared here often.

Of great interest is the appearance of bears at the extreme
east of Siberia, far from the seacoast, often at a distance of hun-
dreds of kilometers from the ocean. These cases are, apparently,
connected with animals wandering away from the ice to the Anadyr’
and Koryat shores. Animals, drifting far to the south, after thawing
of the ice, return to the north by a direct route, 1.e., through the
mainland. In a series of cases, the movement from the south to the
north could be traced successfully and, apparently, the majority of
movements were thus, although there are known cases where
movement was towards Okhotsk Sea; 1.е., to the south. Animals
were encountered in a series of places along the Anadyr’ from its
mouth to Eropol (Utesiki, Ust’-Belaya, П’тиу’уа), Pokul’nya range
(east of the upper Anadyr’), in the upper Penzhina and in Penzhina
region, on the Chernaya river south of Markov on Anadyr’, be-
tween the upper Kanchalan and Tashaina (south of the lower course
of the Anadyr’) and, lastly, in the upper Malyi Anyui.*’ All these
points lie directly to the north of the region of vagrant bears wan-
dering to the Anadyr’ and Koryat coasts. It is possible that some
bears move from the Okhotsk Sea through the Penzhinsk Gulf.

It is natural that in many cases bears are able to penetrate, and
do penetrate, deep into the mainland, regardless of the indicated
type of migration—for example along the Anadyr’, to the south of
Chaunsk Gulf, etc. The extreme northeastern part of the Chukotsk
Peninsula from the Arctic coast (Kolyuchinsk inlet and eastward)
to the Bering Sea, the animals pass overland, apparently even
often”.

Range according to data of Pallas, 1776, 1811; Brandt, 1865; Middendorf,
1876; Pleske, 1887; Iokhel’son, 1898; Zhitkov, 1901, 1904; Buturlin, 1913;
Sokol’nikov, 1927; Ognev, 1931; Adlerberg, 1935; Heptner, 1936; Portenko, 1941;
Pedersen, 1945, 1957; Chapskii, 1946; Averin, 1948 and a series of other authors,
and some unpublished data of S.M. Uspenskii.

““The assumption that the described movement of bears through the mainland
in northeast Siberia might itself be a track of a former transgressions of the sea,
has no scientific foundation.

‘(GaujdaH “D'A) э8ие1 .sivaq ay) jo

AJLPUNOG OY} YIM SPIOUIOD JOU $э0р 31 элэЦи 991 Зицео} Jo Алерипод UJoYJNOS 9Y4}—oUl] иэх014 ay) ‘ззиэшолош BuO] АПеоэ@5э

JO soovjd [елэлэ$ эзвотрит SJOP [eNPIAIPU] ‘5иот5$э185иец 3104$ A[OANL[OI рие SUOIeIZIW [eULIOU JO ио18эл oy) SuIpNyouI эЗиел
э4} JO 1эрлод илэцупо$ 9Y} зэзеэтриг oUT] PI]OS ‘sddiyg 5пилимош (sojoavjoY]) 5п$4[) ‘

Е ЕР

Jvaq 1804 ay} Jo эЗиет 5219946 ‘991 “314 Sor

693
Geographic Range outside the Soviet Union

In the Bering Sea, the range occupies its northern parts (Saint
Lawrence, Saint Matthew, and Hall islands); on the mainland, the
southern boundary passes along the northeastern and northern shore
of the continent approximately from the mouth of the Yukon to the
northern parts of Labrador. Transgressions, frequently together with
floating ice, reach the Pribilof islands, and in the form of special
rarities may perhaps reach the Aleutian Islands, penetrate far to the
south along Hudson Bay, to its extreme southern part (mouth of
Moose river; about 50°45’ N. Lat.). Together with floating ice,
transgressions of the polar bear may penetrate far to the south
along the eastern coast of Labrador—as far as northern Newfound-
land and deep into the Gulf of Saint Lawrence (south; 49° N. Lat.).

The southern border of the range crosses over the floating ice
of Davis Strait, to Greenland and the ice surrounding it, and with
it, encircling the southern extremity of the island and to the ice of
the Greenland and Norwegian seas, including Jan Mayen,
Spitsbergen, and Medvezhii islands. Individual transgressions to the
south reach the northern shore of the Iceland, Varanger fiord and
North Cape (in the past). From this above-mentioned line, the range
extends far to the north—in the same way as in the eastern sector
of the Arctic. In Alaska, transgressions of bears are known deep
into the continent, for 75 miles.

Within the range of the polar bear, differences in the distribu-
tion of places of reproduction (birth of young), winter and summer
migrations, and the regions of seasonal distribution of the animals
of various biological characters (mature males and barren females,
pregnant females, etc.) are very specific. Only in the last decade
(S.M. Uspenskii) were numerical irregularities in the distribution of
the animal and their reasons clarified. The greatest number of polar
bears are, apparently, connected with the so-called “Arctic ring of
life”. It may be represented as a gigantic belt covering the central
Arctic southward and is associated with the 200-meter isobath.
Numerous, usually, large polynyas are associated with this ring, for
example the Velikaya [Great] Siberian polynya—the most produc-
tive part of the high Arctic, in particular very rich in ringed seal
[Phoca hispida}—the main prey of bear. The life and range of the
polar bear are associated, to a significant extent, with this “ring”
(V.H.).

467

694
Geographic Variation

Geographic variation in the polar bear, which from time immemorial
had been considered as one species, was first subjected to analysis
in the 1900’s* by Knottnerus-Meyer (1908). The level of work,
following the worst traditions of Matchie, is astonishingly low—the
author admits 6 species of polar bear, with 3 species for only East
Greenland and Spitsbergen, described by him. The baselessness of
this viewpoint is evident and was shown by several authors (Miller,
1912; Ognev, 1931; Birulya, 1932).

Geographic variation in the polar bear is insignificant. This is,
apparently, explained by two factors—the comparatively small area
of occurrence of the species*' and the considerable exchange of
individuals between different populations. If females are, to some
degree, restricted to definite sections of the Arctic, males roam
very widely throughout the whole year, and violate the isolation of
individual populations, which are generally relative under the geo-
graphical conditions of the Arctic. These wanderings pass both
through the highest polar latitudes, and also, chiefly through the
lower latitudes, apparently, where areas of free water (polynyas) or
broken ice are found throughout the whole year (“Arctic ring of
life’, see above).

There is an idea (Pedersen, 1945, 1956) that, in general, there
1$ an uninterrupted regular movement of roaming bears in a more
or less defined route from east to west around the whole Arctic in
lower latitudes. This leads to total panmixia and completely ex-
cludes the possibility of formation of geographic races. This
conception of migration and its routes, at least for our coastal
sectors and central regions of the Arctic, apparently, requires some
corrections. The point is that the movement of animals from east
to west, which actually takes place, is determined not by any sort
of active “over-flow” of the population by a clock-like pointer, but
by the sum of movements (drift) of the Arctic ice which is, in the
end carried to the Atlantic Ocean between Greenland and
Spitsbergen. However, this “carousel” of the mass of Arctic ice

*In Russian original, “900-kh’—Sci. Ed.

“‘Conceptions about the dimensions of the range of the described species are
usually exaggerated, since Mercator projection maps are most often used. Only
polar or equal-area projections give a correct presentation.

695

is not so simple as it might seem from this very general discussion
of the question. Nevertheless, the main idea deserves attention.

In spite of all, one must never lose sight of the fact that adult
females are relatively sedentary—in winter, they are attached to
their dens and places of reproduction (see above), and in summer
they cannot, with their cubs, roam so widely as males. Their behav-
ior to a certain extent neutralizes the wide movement of males.

It must also be taken into account that, as far as geographic
differences are only manifested in general dimensions, the intensity
of hunting apparently affects the characters of individual popula-
tions. Where animals are intensively pursued, the percentage of
large individuals is, naturally, low. Therefore, the racial unity of the
species is entirely probable. Nevertheless, to completely refute the
known tendencies of geographic (population) variations and to as-
_cribe the differences noted among separate groups on account of
interference by humans is at the present moment still premature.
That itself serves to indicate the presence of some sort of isolation
of populations.

In our country, included in the borders of our sector of the
Arctic, 1.e., between approximately the 30° E. Long. and 170° W.
Long. at least in the lower latitudes, two forms may be noted until
there is a more thorough analysis of the whole question.

1. European polar bear. U. (Th.) т. maritimus Phipps, 1774
(Synonyms polaris, jenaensis, spitzbergenensis, eogroenlandicus,
? marinus).

Dimensions relatively small.

Body length of males in Franz Josef Land (48), 200-М 213—252
cm (most frequently between 200 and 210 cm); females (64) is
165—M 195—236 cm ) most frequently between 190 and 200 cm).”
Body length of males from eastern Greenland (25), 199-M 221.4—

“According to materials of Tsalkin (1936). The series of categories given by
this author contain classes from 110 to 260 cm; i.e., they include young of the
first year; averages are calculated according to this whole material (72 males and
77 females). Naturally, they do not reflect dimensions of animals of this race. Here,
the material is recalculated, and for adults only, males starting with class 180—190
cm and females of class 160—170 were taken. For measurement of separate indi-
viduals, the midpoint of the class (185, 195 and so on) was taken.

468

696

229 cm; females (22), 160-М 169—189 cm.* Animals of this form
from northwestern Taimyr usually have a body length not more
than 230 cm (Heptner, 1936).

Maximum length of male skull, 353-М 355—377.5 mm; female,
334-335 mm; condylobasal length of male skull, 359-372 mm; zygo-
matic width of male, 187.5—M 223—241 mm, female, 194—198 mm;
length of upper molars of males, 56—60 mm (Ognev, 1931).

Weight of adult males from eastern Greenland (normal nutri-
tion) fluctuates between 400-450 kg, and particularly well-nourished
males weigh up to 500 kg. Well-fed females have a weight of 350—
380 kg (Pedersen, 1945).

Found in Franz Josef Land, Novaya Zemlya, the European
coastline (in the past), Kara Sea, western coast of Taimyr (here is
also the following form; see below) and corresponding sections of
floating ice of the Arctic Ocean. ;

Outside the USSR—in Spitsbergen, eastern Greenland and the
ice of the corresponding parts of the Arctic Ocean (for range
boundaries of this form, see below).

For characteristics of this race, it-is significant that very large
specimens in Franz Josef Land and Spitsbergen occur extremely
rarely—of 338 animals taken by different expeditions, only 2 were
larger than 240 cm—one had a body length of 241 cm, and a “huge
old male”, 252 cm. There is information about a specimen from
Franz Josef Land (the method of measuring was not indicated) at
288."

2. Siberian polar bear. U. (Th.) m. marinus Pallas, 1776.

‘According to materials of Pedersen (1945). With respect to Greenland
animals, the size (length) of the body is taken without the tail, as also in relation
to specimens obtained from Franz Josef Land. Therefore, the means given differ
from those given by Pederson (in the data given by him, the average length of the
body is given including the tail and, apparently, even with the end hairs).

“According to materials of Tsalkin (1936). Information about an animal at
370 cm is not believable (Ivanov, cited by Tsalkin, 1936). Data of Pedersen
completely contradict the assertion of Payer that Greenland bears often have a
body length of more than 300 cm (cited by Tsalkin, 1936). The data of Pedersen
(1945) show that the idea concerning the large measurements of Greenland bears,
compared to those obtained from Franz Josef Land, is incorrect, at least with
respect to eastern Greenland. Their measurements are identical.

Individual, particularly large animals sometimes captured in eastern Greenland,
Spitsbergen and Franz Josef Land are probably individuals of the Siberian race
(mostly males) wandering from the east.

469

697

Dimensions larger.

Body length usually between 200 and 250 cm, but often nearer
to latter figure. Separate individuals sometimes larger and attain
300 cm (without tail) and even a few centimeters more. Weight is
between 400 and 700 kg; separate, particularly large, individuals
reach 800 and even approach 1000 kg.*

Skull larger. Greatest length of male skulls, 373-М 386—412
mm; females, 324-М 340-360.5 mm; condylobasal length of male
skulls, 363—М 375—388 mm, females, 327.5—338.5 mm; zygomatic
width of males, 217—M 239-274.5 mm, females, 194—199 mm; length
of upper molars and last premolar of males, 57.6-М 61.7—64.0 mm,
females, 50—61 mm (Ognev, 1931).

Zygomatic width is relatively larger than in U. (Т.) т. maritimus,
and zygomatic arches themselves somewhat more massive, frontal
area abruptly ascends upwards, nasal region relatively longer, lower
jaw more massive, and toothrows slightly longer.

Found on Siberian coast, islands and floating ice from the lon-
gitude of western Taimyr to the Bering Strait.

Outside the USSR. Probably part of the American Arctic.

There exists vagueness concerning the ranges of both forms.
Bears of the western Siberian northward (Ognev, 1931; Birulya,
1932) are usually assigned to the Siberian (eastern) form marinus,
it sometimes being noted that animals of Novaya Zemlya possess
features of a transitional character between this form and western
maritimus. Together with this, there are data (Heptner, 1936) pointing
to the fact that animals of West Siberia eastward to Taimyr are
relatively smaller and they together with Novaya Zemlya animals
are more correctly assigned to U. (Th.) т. maritimus. Thus, in
western Taimyr, in the area between Dikson and the mouth of the
Pyasina lived a small bear (or lived in the 20’s), which apparently

“On the Komsomolskaya Pravda islands (Samuil; southeast of Cape
Chelyuskin, at approximately 77° N. Lat. and 106° E. Long.), on 25 July 1935,
an old male was killed with a body length of 285 cm (without tail) and weigh of
about | metric ton (Shereshevskii, Petryaev, 1949).

Skull dimensions of this animal (Zoological Museum, Moscow University)
slightly exceed even those of the previous animals; its greatest length—413 mm;
condylobasal length—400 mm; zygomatic width—257.5 mm.

On the coast of northwestern Taimyr between Dikson island and the mouth
of the Pyasina, an even larger bear was killed in the 20’s, whose body length was
302 cm (without tail; Heptner, 1936).

698

belonged to the western form (“shore bear” of the native hunters
with body length not more than 230 cm). In winter here, there
appears the large form (“ocean bear”) apparently the Siberian
form И. (Г.) т. marinus. The animals immigrate from the north-
east.

Here there is a quite clear picture of the overlapping of ranges
(for details, see Heptner, 1936). Therefore, the boundaries of the
ranges of the two forms is not, apparently, found in the region of
Novaya Zemlya, but in northwestern Taimyr. Precisely because of
this limit, it follows that the nominal form must be accepted [in
Novaya Zemlya], in spite of prevailing opinion. This is in full agree-
ment with the fact, established by earlier authors, that the largest
bears are encountered in Taimyr and eastward.

Generally speaking, the characters of both forms cannot be
considered established and their independence is often doubted.
Thus, Birulya (1932) saw that the difference between them lies
only in the relative width of the zygomatic arches, and in the Sibe-
rian form, the wider and deeper frontal depression and more strongly
developed processes.

Note. Pallas (1776, p. 691) described not a race, but a species
of polar bear. The description was done by comparison with the
brown bear, because before him, the polar bear was poorly known
and unsatisfactorily described. The distribution was described as on
the peninsulas and islands, and also on the pack ice of the northern
Arctic Ocean, from which it nowhere moved away (“habitat in
promontoriis insulis atque glacie fluctuante Oceani Hyperborei, a
que nusquam recedit”). The first-hand material for description con-
sisted of a young animal from the southern part of the Ob’ Gulf,
which was transported alive to Krasnoyarsk, where Pallas wintered
with his student and fellow-traveler Vasilii Zuev (Pallas, 1776, p.
38). Therefore, in the latest literature, the type locality of the form
marinus is correctly considered the Ob’ Gulf.

At the same time, no one has made a direct comparison be-
tween Ob’ bears (they have been absent for a long time) and the
Spitsbergen bears. At that time, all authors, noting the large size of
Siberian animals, referred [them] to animals from the shores of
Middle and Eastern Siberia and the Far East. The above mentioned
views show that bears of the western and southern parts of the
Kara Sea do not differ from those of Spitsbergen. For the Ob’
animals, it is certain, according to the query of Zuev, that Pallas
(1776) referred to quite large measurements, though less than those

699

of eastern [bears}—up to 227—260 cm (converted from the Parisian
feet which Pallas used—‘septem et octo saepe pedes adultorum
magnitudo”).

Therefore, the name marinus is formally included in the syn-
onyms of the nominal form, but the eastern bears must receive a
new name. Until variation in our bears has been thoroughly revised,
it is also undesirable, and it is proposed to retain the name marinus
but to consider its type locality not Ob’ Gulf but the Laptev Sea,
where the large form undoubtedly occurs.

At present, it is difficult to give an opinion about the forms
living outside the boundaries of our country, except for eastern
Greenland (see above). All three subspecies described for the
American Arctic—groenlandicus Birula, 1932; ungavensis
Knottnerus-Meyer, 1908 and /abradorensis Knottnerus-Meyer,
1908, were described from places lying near one another (western
Greenland, Labrador and Ungava Gulf respectively), and are, ap-
parently, identical. American authors usually assign bears of their
Arctic areas to the nominal form (Hall and Kelson, 1959), although

Fig. 167. Dresh-Kheb mountains—tegion of greatest concentration of bear
dens on Wrangel’ Island. Beginning of April 1964. Photograph. by
S.M. Uspenskii.

470

700

there are still some authors who even now recognize the “spe-
cies” Jabradorensis (Anderson, 1946) and other “species” after
Knottnerus-Meyer (Miller and Kellog, 1955). In the American Arctic,
very large animals occur, but not inferior to North Siberian bears
(condylobasal length of skull, 391 mm; Miller, 1912) (V.H.).

Biology

Population. Over a wide area of the Arctic Ocean the polar bear
population is distributed unevenly. Their greatest concentration is
observed in the region of confluence of the variously moving masses
of sea ice. This region encircles the central Arctic, approximately
at the level of the 200 m isobath (“Arctic ring of life”; p. 693).
The second region of concentration is at the edge of drifting ice,
mainly near the Atlantic sector (ОзрепзКи, 1961). The population of
polar bears is not stable: the animals periodically shift in different
directions. Moreover, passive displacement, conditioned by continu-
ous drift of sea ice from east to west in clockwise direction with
a speed of about 2.3 miles per day, is of substantial importance
(Pedersen, 1945; Uspenskii, 1961).

During the last 40—50 years the number of polar bears was
strongly reduced as a result of their destruction. At the present
time, according to various data (aerial count, den count, etc.), its
total number was determined as 5—6 thousand head (Uspenskii,
1961). In the Soviet sector of the Arctic, their number hardly ex-
ceeds 2—3 thousand head.

Habitat. The polar bear is a marine animal. The main place of
occurrence is floating ice of the Arctic Ocean, and to a lesser
degree, the coastal zone of the sea-coast and its islands. The polar
bear rarely moves on to the tundra, and usually not far but, some-
times, penetrates to the tree-line (along the Kolyma, 10 km above
the mouth of the Omolon), usually in the period preceding their
hibernation in dens, and in spring, when thaw-water lakes form on
the ice, and cracks are still absent. On the tundra at that time it
restricts itself to the valleys of creeks in ravines.

The existence of polar bears is closely connected with fissures,
cracks, edges of ice floes and fast shore ice along the coast, be-
cause it is only here that they are able to catch their principal
food—seals (ringed and bearded seals;* Rutilevskii, Popov and

*Phoca hispida, Erignathus barbatus—Sci. Ed.

—

701

Shastin, 1939). Therefore, the southern boundary of the species
range in Asia is limited to the northern coast of the continent, and
in the Atlantic Ocean—the southern boundary of floating ice, and
changes depending upon the position of the floe ice (Birulya, 1932).
In Alaska, 1 bear traversed 32 square miles, with a range of 21 to
57 miles (observation from the air, Scott and others, 1959).

Food. The main food of bears is ringed seal, to a lesser degree,
bearded seal. In the stomachs of 145 animals killed in Franz Josef
Land (Tsalkin, 1936), ringed seal was found in 67% of stomachs,
walrus—in 14.4%, bearded seal—in 5%, vegetal mass—in 4% and
birds—in 0.6%. Polar bears eat the carcasses of hunter-killed wal-
rus (L. Popov, 1939), but themselves do not attack them because
the adult walrus is a prey beyond its strength and the young ones
are defended by the adults (Leonov, 1953). When bears and wal-
ruses meet, they pay no attention to each other (Rutilevskii, 1939).
The bear hunts seals from ambush, waiting for them at their holes
or cutting off their path to the water. The polar bear feeds most of
all on ringed seals, mainly by virtue of their forming, in definite
places, considerable accumulations (Pedersen, 1945). Ringed seals
are subject to attack when resting or sleeping near air holes in ice.
The most intensive hunting of ringed seals by polar bears occurs in
spring and the beginning of summer, and at the time of molting,
when the animals are less cautious; they employ a stalk using
concealment (ice-hummocks) and hiding near the hole. Approach-
ing to 4-5 meters, the polar bear makes a big jump. During an
ambush, the bear waits until the head of the seal does not appear
from the hole above the water* then, it strikes the seal’s head with
its paw and draws it to the edge of the ice-floe. The white bears
very rarely encounter migrating seals—harp [Phoca groenlandica]
and hooded [Cystophora cristata].

It usually eats the fat and skin. Only a hungry animal also eats
the seal meat, leaving the bones. At the age of 6 months the cub-
bear eats 2.5 kg of seal fat; the adult—6—8 kg (Rutilevskii, 1939),
rarely more—up to 20 kg (Leonov, 1953). There is an opinion about
its capacity to eat even 71 kg of walrus meat (Naumov and Lavrov,
1948), but this is an exaggeration. Polar bears do not themselves
catch fish (Rutilevskii, 1939; Leonov, 1953). Hungry bears eat not
only carcasses of seals and walruses, but even salted cod, carrion,

*Meaning unclear; perhaps the negative statement is a Japsus—Sci. Ed.

472

702

and also molting and nestling birds which fall from nests, and marine
flotsam. There are known cases of their eating reindeer and Arctic
foxes (Esipov and Pinegin, 1933). This accidental food occurs pri-
marily in that period of hunger for the polar bear, when marine
animals are not available. Depredation on stores of foodstuffs of
the polar expeditions by polar bears is a common phenomenon.

Home range. Polar bears are extremely mobile, roaming widely,
and have no permanent home range boundaries.

Burrows and shelters. The polar bear belongs among that
group of animals that do not have permanent shelters. As they
wander about the pack ice, they rest by lying among ice-hum-
mocks. Obtaining its food within the limits of the coastal zone, the
bear prefers to move to the sea-ice to lie down (L. Popov, 1939).

In winter, females, and in some cases males, construct dens in
snow drifts. In Novaya Zemlya, such dens sometimes have 2—3
sections, including a “toilet”. In four investigated dens, only one
chamber was found (Esipov and Pinegin, 1933). After construction
of the den it usually covers them with snow, forming a large snow
drift, in which a small opening sometimes remains, serving for breath-
ing, but usually it is also blocked.

Dens of young bears are distinguished by careless construction
and site selection. The den is usually oval inside, its length is 160—
180 cm, and height 100—120 cm. Dimensions of the den, where the
she-bear overwinters with cubs, reaches 260 cm. The length of the
passage leading to the den chamber is 2.0—2.5 т, and even 3.0 тт,
rarely—l10 т, and the diameter of the entrance is 60—70 cm
(Pedersen, 1945).

On Pinegin Island, a den was found with a tight, dim corridor
that was about 6 m long; the chamber was as high as a person and
it had a diameter of about 3 m (Pinegin, 1933). On the pack ice,
there are no dens; bears build them on the shores of the mainland
and islands, frequently under over-hanging shorelines, among stones,
rocks and so on, where large snow drifts are formed.

Dens are distributed primarily on the Arctic Ocean islands, but
in part along the sea coast. In the eastern Arctic, this is mainly in
the Franz Josef archipelago, Wrangel’ Island, and also the New
Siberian islands. It is assumed that there are 1000—1500 dens in the
entire Arctic. If we calculate that each is occupied by a pregnant
female (with which not all investigators agree) and that the latter
comprise about 20% of population, the total number in the popula-

703

tion is determined to be about 5000—6000 head (Uspenskii, 1961).
In spring, when the litters have not yet emerged, perhaps a maxi-
mum of 8000 may be estimated.

Daily activity and behavior. The peculiarities of the light
regime of the arctic do not facilitate the performance of a distinct
rhythm of daily activity in the polar bear. Its life is closely con-
nected with t1e sea and the pack ice; it is a true marine animal; it
can swim ana dive perfectly. The thick layer of fat and the dense

Fig. 168. Entrance of den recently abandoned by she-bear with cubs.
Wrangel’ Island. End of March, 1964. Photograph by S.M. Uspenskii.

473

704

fur permit the bear to remain in cold water for an extended time,
and its strength and endurance in swimming are so great that it can
swim for 10—20 and even 35 km. In the region of Wrangel’ Island,
the polar bear was observed in the open sea, 100 km from the edge
of ice (N.N. Kondakov).

It swims at a speed of 5.5—6.5 km per hour. It can stay under
water for 2 minutes. In and near water, it feels more confident and
bold than on land, where it is not distinguished by its activity. It does
not run qui
that of males while fattening is up to 5.0 km/h. It trots 8—10— 12 km/
h, and in a gallop, up to 15 km/h, but such speed is possible only
for a short distance. During quick movement, the polar bear quickly
becomes exhausted. A human can tire a large polar bear after a
distance of 10 to 15 km, because even after 5 km, its speed begins
to decrease. However, in even large massive ice-hummocks, it
maneuvers amazingly adroitly, and easily escapes from dogs and
human. In so doing, the bear can jump across ridges of 1.5 to 2.0
m high and jump to a height [width ?] of 4-6 т (Rutilevskii, 1939).

The sense organs of the polar bear are well developed. It can
see seal carcasses on the ice at 2—3 km. It can scent a strong odor
(hot fat) at 6-7 km (with a fair wind). It hears the approach of a
human at 200 т (Rutilevskii, 1939). Where it is not pursued, it is
bold and self-confident and exhibits great curiosity towards un-
known objects. Polar bears attack man extremely rarely, even when
hungry; an unexpected meeting with a female bear with her cubs
is somewhat dangerous.

They tolerate a temperature of 10—15°C вон, and seek shade
(Kost’ yan, 1954).

Hibernation and winter sleep. Information about the winter
sleep of polar bears is quite contradictory. There are some indica-
tions that in Novaya Zemlya, males are active the entire winter
(Esipov and Pinegin, 1939). This sometimes extends also to barren
females and young bears (Lavrov and Naumov, 1948). The belief
exists that only pregnant females, with few exceptions, retire to
dens. They point out that old and strong bears are active in winter,
whereas the younger hibernate for some time (Heptner, 1932).
Among females the greater part hibernate in winter. A consider-
able portion of animals in the more southern regions, which are
better supplied with food, do not den at all; those which do den,

705

Fig. 169. Polar bear running at a gallop. Wrangel’ Island. Middle of April,
1964. Photograph by S.M. Uspenskii.

hibernate for a short time. Only a small portion of them hibernate
from autumn through the whole winter.

Winter sleep of polar bears is related to food deficiency in
winter, and to difficulty in obtaining it in the dark period of the year.
Under more favorable conditions in the southern part of the range,
where this period is shorter and it often happens that the sea is not
frozen, sleep is shorter and the animals den more rarely. On the
northern coast of Greenland, 90% of the animals den, in the north-
erm part of Baffin Land—S0%, and in southern Greenland—only
30%. On the whole throughout the range, 70-80% of the animals
den (Pedersen, 1945).

Finally, there is information that all polar bears retire to dens,
but for different periods: adult males den for 50—80 days, emerging
from the den on the first days of February; non-pregnant females
retire until 13—19 March, for a period of 115—125 days; females with
cubs of the current year—auntil the first days of February for a
period of about 160 days; and pregnant females retire to dens for
160—170 days, emerging from them at the end of April (Rutilevskii,

474

706

1939). In the region of Cape Chelyuskin, barren and pregnant fe-
males lie down to hibernate in the second half of November, as also
do the cubs of the current year (Rutilevskii, 1939). Adult males
retire later—by 12 December. On the eastern coast of Taimyr, a
male was killed in a den on 9 December 1935 (L. Popov, 1939).
Hibernation of males has been confirmed for the Olekma.

On the Olekma, emergence of bears from the den is noted in
March—April. On Severnaya Zemlya, emergence from the den is
extended, the principal mass emerging in the second half of March
(L. Popov, 1939); on Novaya Zemlya, until the last days of April
and even until the twentieth day of May (Pinegin and Esipov, 1939).
On the whole, the time of retirement of bears coincides with the
beginning of the dark period in the Arctic: in the south—this is the
end, and in the north—the middle of November. The period of
emergence is more prolonged—from the end of February to the
second half of March and not later than the beginning of April. Old
males are the first to den and the first to arouse (Pedersen, 1945).
Delay in time of emergence of she-bears from the den is connected
with delay in birth of the cubs.

Seasonal migrations and transgressions. Polar bears wander
widely within the limits of the Arctic expanses they inhabit. Their
migration is influenced by: 1) condition of the pack and 2) the
distribution of seals (ringed and bearded seals), which depend upon
the sea’s ice cover conditions.

In summer, polar bears everywhere migrate to the north, and
in winter—to the south. By winter, animals entering hibernation are
concentrated on the islands of the Arctic Ocean (Severnaya Zemlya,
Franz Josef Land, Wrangel’ Island and north Novaya Zemlya) and
in the coastal zone. On Bennet Island, in the period of mass migra-
tion of polar bears, tens, and up to 14 bears at one time may be
seen daily passing from regions of solid ice to open water (Uspenski,
1961). On the eastern coast of Taimyr, polar bears arrive from the
north, northwest and northeast in winter (L. Popov, 1939).

On Severnaya Zemlya, from May, when the ice is solid, they
move out far from land from one [open water] lead to another,
making a whole “bear highway” (Ushakov, 1951). If there is no
[open] water, they move to the promontories and spits of land, where
cracks and small leads, which attract ringed seals, are often found.
With appearance of solid ice, they frequently move to fast shore
ice, retreating southward.

475

707

Polar bears roam widely in the region of Cape Chelyuskin; the
main direction of their movements is from east to west. There are
also local and seasonal movements from north to south and back
again. Those leaving from dens in spring to the west of Taimyr go
to the east and north. Near the coast, in June—July, they migrate
mainly to the west, where marine animals are concentrated
(Rutilevskii, 1939). Animals appear in the northeastern part of the
Kara Sea in the first half of winter, migrating from nearby regions.
In the middle of winter large bears appear, arriving from the remote
parts of the Arctic Ocean. With the approach of spring, they gradu-
ally migrate again to the northeast along the coastal fast shore ice.
Bears already adjacent to places remain for a longer time and the
last of them depart in June (Heptner, 1932).

In Spitsbergen, polar bears migrate following the southern limits
of the pack ice, near which seals are found; they constantly move
between Spitsbergen and Novaya Zemlya, appearing periodically
on Medvezhii [Bear] Island. To the north, they go as far as the
region of the pole, occurring both on solid and on broken ice (Birulya,
1932).

Cases are known of drifting of polar bears on fields of pack ice
far to the south—to St. Matthew Island in the Bering Sea (Sokol’nikov,
1927), and the Okhotsk coast (G.D. Dul’keit). During migrations,
they sometimes accomplish long traverses (10 km and more) of
tundra (Sokol ’nikov, 1927) (for details, see section on “Geographic
Distribution’).

Reproduction. On Novaya Zemlya, rut commences in March
(Esipov and Pinegin, 1933), but there is some indication (Dubrovskii,
1937) that it is prolonged here from April to August. On Severnaya
Zemlya, it is observed from the middle of June to July (L. Popov,
1939). In the region of Cape Chelyuskin, the males and females
were observed to stay together from 2 July to 27 August (Rutilevskii,
1939).

Outside the boundaries of the USSR, the breeding period* oc-
curs from the end of March or beginning of April to the end of that
month. In the absence of conception, there is a second, late, estrus
in May. Estrus in females lasts 3 weeks and is usually repeated
every 3 years (Pedersen, 1945)—according to other data, every

*Misspelled “perid” in Russian originatSci. Ed.

708

other year, but in the absence of conception, even earlier (Kost’ yan,
1954).

Usually, 3 to 4 males go after a female who comes into estrus.
Therefore, it is supposed that the ratio of sexes equals 3 to 1;
however, up to 7 males, moving at considerable distances from
each other, have been observed with one female (Pedersen, 1945).
If estrus in females actually occurs every 3 years, then approxi-
mately 1/3 of the adult females are found in estrus each year, and
the presence of several males near the female is fully understand-
able.

In the Leningrad Zoo, the period of rut proceeds annually from
15 March until the end of April (Kost’yan, 1954). If copulation
takes place in July or August, the female remains barren. Copula-
tion in the period of rut is repeated many times. The duration of
pregnancy ranges between limits of 230—250 days, 1.е., about 8 months.

The behavior of the pregnant females changes noticeably by at
least October, when they turn towards the nearest land. This is
especially noted where bears are far away in the open sea. Besides
large mountainous islands, pregnant females occupy dens on small
“bear” islands. Such females often build snow-burrow dens not on
the coast, but several kilometers from it. The den is constructed
from the end of October—-beginning of November, and is occupied
from the middle of November, when embryos development begins
after the latent phase.* Embryos were never found in females
killed outside of dens (Pedersen, 1945).

On Novaya Zemlya, according to various data, birth of the
young takes place in January—April (Dubrovski, 1937), [or] in De-
cember—January; on Krestova Bay, in March—April (Esipov and
Pinegin, 1939), and beyond the borders of the USSR (Pedersen,
1945), during January. In the region of Cape Chelyuskin, parturition

15 observed in January—February, sometimes later (Rutilevskii, 1939).
On Severnaya Zemlya, birth of young occurs in the middle of June
and even in July (Popov, 1939). Although in the polar bear, as in
species with a latent stage in embryonic development, the time of
birth may be shifted significantly due to variation in the duration of
this stage, claims of June—July are undoubtedly in error. In general,
one may state that the time of birth occurs in the middle and second

*Delayed implantation—Sci. Ed.

ee еее.

476

709

half of winter. In the Leningrad Zoo, birth of young in polar bears
took place from 10 November to 28 December (13 cases).

The number of the young bears in a litter equaled two in 11
cases, and one in 2 cases (Kost’yan, 1954). Under natural condi-
tions, the number of young varies from one to four. Of 27 cases
(different authors), the litter contained 2 cubs in 16 cases, and in
10 cases—one, and in one—4 (Pedersen, 1945). The cubs remain
with the female bear about 1—1/2 years.

Growth, development and molt. Bear-cubs are born helpless
and blind, with closed ear openings. Wool is sparse and short, and
skin pigment is absent. On the 3rd day after birth, body length is
28—32 cm, average weight—about 755 g (650—840 g). On the 26th
day, body length increases inconsiderably—to 35—38 cm, but weight
grows to 1125—1275 g. Pigmentation [of skin] is completed by the
end of the first month. Eyes open on the 30th—31st days; at that
time the ear openings also appear. They begin to crawl at the age
of 45 days. The first signs of the appearance of sense of smell are

Fig. 170. Bear-cubs which have just emerged from the den. Wrangel
Island. 15 April 1964. Photograph by S.M. Uspenskii.

476

710

observed at the age of 50 days. Teeth are cut at the end of the 2nd
month. At this time, the cubs regularly crawl out of the den, but are
capable of short walks with their mother not earlier than 3 months
after birth. They feed on the mother’s milk for 6-8 months (Kost’ yan,
1954). In individual cases, they suckle the mother up to 15—18
months (Rutilevskii, 1939).

At the age of 60 days, males weigh about 7 kg, females about
6 kg, with body length of the former 58—60 cm, of the latter, 54—58
cme

At an age of 4 months, cubs have attained a weight of 22—23
kg; at 6 months of age with length of about 110 сп-—40—43 kg, and
at 8 months—58—62 kg. Body length from the age of 6 to 9 months
increases by 6—7 cm per month. At the time of winter sleep, growth
almost ceases. They begin to swim at the age of six months.
Replacement of teeth by permanent ones is completed by the 10th—
11th month; at the conclusion canines are replaced by October
(Rutilevskii, 1939). Sexual maturity is attained in zoos at 5 years
(Kost’yan, 1954). In Greenland, the females participate in repro-
duction at 4 years, and the males—8 years (Pedersen, 1954).

In contrast to males, females with cubs restrict themselves to
coastal areas, and do not move far, either into the tundra or the sea.
Yearling cubs abandon by the mother subsequently stay together
for about 7 months.

The she-bear leaves the den at the end of March—beginning of
April, rarely earlier, when the cubs are 8 to 9 weeks old. Some-
times for several days thereafter, she remains near the den, and
then goes out to the coast near sea ice. At first, she sometimes
leaves the cubs in a den. While hunting seals, the she-bear leaves
the bear-cubs in a secure place, but never on the mainland, or else
they accompany her.

Adult males are a danger to the cubs, and she-bears avoid them
in every way. They also, though very rarely, move to islands and
the sea coast. Usually, the cubs spend the first winter in a den with
the she-bear, who leaves them at the end of their second year of
life, when they are equal to her in size. However, the cubs some-
times spend a second winter with the she-bear and go with her in
spring to hunt seals (Pedersen, 1945).

Enemies, diseases, parasites, mortality, competitors, and
population dynamics. Except for humans, the polar bear has no
enemies. In the Arctic, the polar bear also has no competitors for

711

477 their main food—seals. On land, dogs and wolves are enemies of
the polar bear (Pedersen, 1945). In water, it is walruses which it
fears most of all, and the killer whale (Orcinus orca).

Diseases which markedly influence changes in numbers are not
known.

There are no data on mortality, but it may be assumed that cubs
naturally die from starvation, being carried away on pack ice, and
others like it in magnitude. Death of adult bears is determined by
their economic use by humans, starvation and other accidental causes
(carried away on pack ice, etc.). Natural mortality among adults is
probably not high. During the last decades, trichinellosis was very
widely spread among polar bears (S.M. Uspenskii).

Field characteristics. It is impossible to confuse the polar bear
with any other animal in the polar regions of the Arctic.

Its tracks are distinguished by the marks of the powerful claws
on the forelimbs. In contrast to the brown and white-chested

Fig. 171. Tracks of a she-bear and cubs traveling from Wrangel’ Island to
the ice. Middle of April, 1964. Photograph S.M. Uspenskii.

478

(Himalayan) bears, the lower surface of the feet of the polar bear
is covered with wool. Therefore, the large, broad and flat track of
its hind limbs are more similar to those of the track of a human foot
shod in fur shoes, than to the print of his bare foot, which is
characteristic for the brown bear (P.Yu.).

Practical Significance

The skin of the polar bear has limited use, and is principally used
for decorative purposes (carpets). There is no exact information on
the number of polar bears obtained in the Soviet sector of the
Arctic before the prohibition of their taking in 1956, although it is
calculated to be in the hundreds of animals. The yearly take of
polar bears in the entire Arctic in 1957—1958 was determined to be
950—1400 individuals, of these 100—200 in Alaska 400—500 ш Canada,
150—200 in Greenland, 150—300 in Norway, and in the USSR, pre-
sumably 150—200 (Scott et al., 1959). The main significance of this
animal lies in the fact that it serves as an emergency stock of meat
and fat for the members of Arctic expeditions wintering in polar
stations and so on. Only in these events and in cases of direct
danger to life, is the killing of polar bears permitted.

Hunting takes place on the ice by rifled weapons, ambush,
stalking and driving, sometimes with the help of dogs. There also
exists another method of hunting polar bear from specially con-
structed, baited blinds.

As early as 1954, foreign organizations for nature protection
have raised the question of the necessity of complete protection of
this animal in all sectors of the Arctic, and in this sense have
applied measures.

In the USSR, by decree of the government, hunting of polar
bear has been prohibited since 1954. Only in emergency conditions
is it permitted to expeditions and overwintering people at polar
radio-stations. Hunting of polar bear (and snow goose) on Wrangel’
Island, the main concentration area of the dens of pregnant she-
bears was declared illegal. All these measures have helped to prevent
the extermination of the polar bear; however, convincing data about
growth of their numbers are still absent. The fate of this animal still
fills us with apprehension, especially as a certain number of bears
are known to be killed yearly under the pretence of self-defense
and for capturing cubs for zoos.

479

713

Counting from the air in Alaska in the late 50’s showed some
increase (1956—1 animal per 148 km route, 1957—1 animal per 56
km, 1958—1 animal per 55 km; Scott et al., 1959), although these
indications may be based upon accidental causes (P.Yu.).

Subgenus of the White-chested Bears

Subgenus Selenarctos Heude 1901
WHITE-CHESTED, BLACK BEAR“
Urusu (Selenarctos) thibetanus G. Cuvier, 1823

1823. Ursus thibetanus G. Cuvier, Rech. ossement fossiles, 4, p.
325. Assam.

1841. Ursus torquatus Wagner, Schreber* Saugethiere, 2, р. 144.
Renaming of thibetanus.

1901. Selenarcios ussuricus Heude, Мет. Hist. Nat. Emp. Chin.,
5, pt. 1, p. 2, pl. 11, fig. 10. Ussuri Territory (V.H.).

Diagnosis

Size moderate. Coloration black, very constant; on chest, a large
white spot. Lower surface of forefeet bare (covered by one con-
tinuous callosity). Facial part of skull relatively short, cranial part
large, broadened; sagittal crest weakly developed, last upper molar
wide and long, considerably larger (almost twice) anterior, last lower
molar small, not narrower posteriorly (У.Н.).

Description

In general appearance, the white-chested bear is similar to the
brown bear, but its build is lighter—the body is not so massive, and
limbs are thinner, taller and slender. In contrast to the polar bear,
the anterior part of the body is more powerful than the

Тре present, very apt, native name for this species in the Far East—white-
chested bear (Bromlei, 1956). The second name is pedantic. In the literature, this
species is called also the Himalayan and Tibetan bear. It is not necessary to retain
this artificial nomenclature, the more so since the latter gives a false impression
about the range of the species.

*Misspelled Shreber in Russian original—Sci. Ed.

714

posterior, and the hind limbs are weaker than the forelimbs.’’ In
motion, the white-chested bear looks as if its hind legs are crossed
with its short legs under its body. The main feature mentioned
about the build of this bear is more pronounced due to the fact that
it has high withers with long hairs, and when moving, its neck and
head hangs down low. The head is relatively small, and quite nar-
row, with an elongated muzzle. The ears are very large and broad,
narrow at base and strongly protrude from the fur, giving the animal
a unique appearance.

The lips and nose are very mobile, and larger than those of the
brown bear. The tail is short and hidden in the fur. The claws are
large, and more abruptly curved as compared to black* bear; claws
of the hind limbs are relatively longer than in the brown bear; and
differ slightly in length from the forelimbs (ratio of length of anterior
to posterior is 1.17: 1.00 against 1.74: 1.00 in the brown bear).
The claw color is black.

The hind sole has 5 well developed bare digital pads (callosi-
ties), the base of the free part of the digits being covered with hair;
the remaining lower surface of the sole is bare, as in the brown
bear. On the forelimbs, not only are the digital pads bare, but also
the entire lower surface of the foot.

Fur is dense, fairly long and fluffy. Hair length on the back
reaches 100—105 mm; on the withers, occiput and neck is even
longer and along the sides of the neck reaches 160 mm. In the
thoracic region and abdomen, the hair is, on the contrary, short and
has an overall length of about 20 mm (in the brown bear, it is very
long in these regions—200—205 mm). In winter coat, underfur is well
developed; in summer, it is absent, and the fur is shorter.

Coloration of the winter coat is very beautiful, brilliantly black,
with unique reddish brown highlights on the head and the thick
underfur, and sometimes on the sacrum. In summer, when the

“’The ratio of the weight of the disarticulated anterior extremity to the weight
of the posterior in the white-chested bear is 1.1 : 1.0, in the brown bear, 1.0: 1.05
(average of 6 specimens; Bromlei, 1956). An individual taken on 5 August 1940,
the weight of the two hind limbs, disarticulated at the trochanteric joint, comprised
9 kg; the two fore limbs together with shoulder blade was 10.6 kg. The forelimbs
are so strong that an animal with broken hind limbs can move on its fore limbs
and even climb a tree with, knots though not high (G.F. Bromlei).

*Meaning not clear; may refer to brown bear, or to American black bear—Sci.
Ed.

480

WIS

Fig. 172. White-chested, or black bear, Ursus (Selenarctos) thibetanus G.
Cuv. Sketch by A.N. Komarov.

undercoat disappears, the coloration is pure black. On the thorax,
a large, almost always sharply outlined light patch is found. Its
outline is quite variable, but it always elongated in a transverse
direction and generally has a half-moon or crescent form, with the
ends of the “crescent” directed forward; sometimes from the
middle part the projection of white coloration is directed back-
wards.

Occasionally, the patch has an oval form, or is divided into two
(a larger part lying above, and a small—below) or even into three.
In rare cases, the patch may be completely absent. In our bears the
patch is most often in the form of an angle (“izhitsei”)*. A white
patch is sometimes situated on the lower lip and chin. The color of
the patch is usually pure white, sometimes with a more or less
ocherous film. There is no sexual difference in color, and the indi-
vidual variation, not counting the white patch, is negligible.

The skull is relatively small but massive, with massive lower
jaw. In contrast to the other two species of the genus, projections

*Name of a V-shaped archaic letter in Cyrillic alphabet—Sci. Ed.

716

and crests of the skull are weakly developed. The sagittal crest is
low and short, even in old individuals, and is only noticeable on the
posterior part of the parietals; it never extends beyond their anterior
edge. Its length does not exceed 19-20% of skull length (in the
brown bear, it may comprise up to 41%). From the sagittal crest,
two arciform flattened ridges (crests) extend forward, reaching the
supraorbital processes. In this manner, the frontal area, which in
other species has a rhomboid form, in the white-chested bear ex-
tends backwards as a long projection to the parietal bones.

The general profile of the skull is a gently sloping arch, and a
frontal projection is not formed. The frontal area is neither im-
pressed nor flattened, its transverse profile being slightly convex.
The supraorbital processes are relatively weak (weaker than in
other bears) and depressed (not elevated). The occipital crest is
well developed, its ventro-lateral part particularly large and broad,
so that the whole posterior part of the skull is broad. The zygomatic
arches are quite weak, but in the posterior part, in accordance with
the size of the braincase, they are widely separated. Their posterior
part, in the region of articulation with the lower jaw, is greatly
flattened and broadened transversely.

One of the main characteristic features of the general skull
structure includes the strong shortening of the facial part of the
skull, and the broad and voluminous posterior expansion of the
braincase. The latter feature is emphasized by the structure of the
occipital crest indicated above. In combination with the weak de-
velopment of the crest and the general sculpture of the skull, and
the almost undefined postorbital constriction of the cranium, this
confers on the skull of the described species an infantile appear-
ance.

The distance between the anterior end of the premaxillary bone
to the line uniting the ends of the supraorbital processes is consid-
erably shorter than the distance from this line the end of the sagittal
crest (or the mid-point of the occipital crest). The distance from the
anterior edge of the orbit to the anterior point of the premaxillary
bone is less than the distance between the ends of the supraorbital
processes. The muzzle is relatively broad, its width above the ca-
nines is equal to the width between the infraorbital foramina or
even exceeds it. In concordance with the general shortening of the
facial part of the skull, the lower jaw is also shortened.

482

717

The toothrows, compared to those of the brown bear, are rela-
tively short (comprising about 40-41% of the total length of the
skull), the canine is strong, and the other teeth are, in relation,
somewhat weaker. On the whole, the dental formula is the same
as that of the brown bear; in particular, the last upper molar is
particularly large. Compared to it [brown bear], however, [this molar]
is slightly smaller—its size is somewhat less than twice the length
of the first molar. The last lower molar is small and is much smaller
than in the brown bear. It has a rounded-rectangular or oval form,
usually without the constriction in the posterior part characteristic
of the brown bear (Fig. 150). The cusps on the surface of the
molars are more weakly developed than in the brown bear. The
first three premolars, which often fall out with age in the brown
bear, are usually retained.

Dimensions are considerably less than in our other bears, and
correspond approximately to the dimensions of small races and the
individual variants of the brown bear (Table 47).

Large males have a size [body length] of about 150 cm;*
occasionally including bears with body length of more than 170 cm,
and very rarely those of 200 cm.* The greatest weight of the bear
is in autumn, especially in years of good food harvest, when they
put on much fat before hibernation.

Table 47. Dimensions of animals (in the flesh) from southern part of
Ussuri territory (Tachin-Chtan range) (Bromlei, 1956)

Specimen Body Tail Ear Chest Height at Weight in
length length length circumfe- withers kg
rence

Males

6 January 168 10 13 134 65 LI?
18 February 153 9 15 74 87 110
9 November 171 11 12 125 86 147
Females

30 October 128 4] 12 102 64 65
24 November 135 7 15 95 82 80

12 Мау 144 10 18 89 79 69

*Contradicts Table 47, where mean body length is 164 cm—Sci. Ed.
“On 3 December 1940, Yu.A. Salmin killed a male in Terneisk region with
a body length of 197 cm (G.F. Bromlei).

718

PPPS
aN \ Te
33) VV. О
a, =a 6 yy > SS . 3
Z\> BRN SAV H R
> =RW S AS
\ 5

ИХ

м -
Е
Wtf Yay se
+

YN,

ИХ
И

Hh)

«а

TM Eis? yt
“ а

227

й
4

72
vente
(К

ААА
(Е
oy
a
—/.
727,
oT ffir
pat p

Tite,
ee
2. Ay i
7 Ат
м

LALLA ER

+ Звмсиииии"
й
SW
57

At “ ;
|) GOS Ry

д ©: AS

АИ $

И”

t х

hye <
и THAN AW

481 Fig. 173. Skull of the white-chested bear, Ursus (Selenarctos) thibetanus
G. Cuv.

483

Greatest length of skull of males (10), 295—M 311.7-328 mm,
condylobasal length of skull of males (10), 271-М 291.6-315 mm, of
females—260—268 mm; zygomatic width of skull of males (9), 185—
М 199.5228 mm, of females—163—173 mm; length of upper toothrow
in males, 110—112 mm, of females—98—100 mm (from materials in
Zoological Museum, Moscow University and the data of Ognev,
1936).

Females are always somewhat smaller than males. Individual
variation is not great, and has a “normal” character. There is
nothing of the kind of variation that occurs in some brown bear
populations. Geographic variation in the size exists, but is generally
small and, in the majority of cases, less than that in the brown bear.
It is not pronounced within the boundaries of our country.

Systematic Position

The white-chested bear must, obviously, be considered a more
specialized form than the brown bear. In this respect, it is analo-
gous to the polar bear, although as regards its own degree of
difference, it stands farther from the brown bear than does the
polar bear.

Moreover, specialization of the white-chested bear has a com-
pletely different, so to say contrary, direction as compared to the
polar bear. As was mentioned above, the polar bear is an entirely
terrestrial* animal, a good swimmer, and a true predator. The white-
chested bear is an herbivore to a greater degree than the brown
[bear], in whose structure may be observed completely evident
features of adaptation to tree-climbing (development of the fore
limbs and claws, short fur on the chest and abdomen etc.). In
Ussuri territory, this species spends up to 15% of its time in trees,
feeds there, and hibernates in hollows. The brown bear can also
climb trees, but only up to the second year of age (Bromlei, 1956).

Biologically, but also morphologically, the white-chested bear
itself represents the beginning of that line of specialization which is
maximally attained in the South-Asian arboreal forms—the sloth
(Melursus ursinus) and Malayan (Helarctos malayanus) bears.

*In the sense that it does not climb trees; as noted above, it is primarily
marine—Sci. Ed.

720

In systematic relationships, however, it apparently has less in
common with the sloth bear; but (attention was not drawn to this
point) its skull has common features with the skull of the black
Malayan bear. This similarity is more obvious when compared with
the skull of a young Malayan bear.

The skull of the adult animal [of Helarctos] differs strongly,
but some of its characteristic peculiarities themselves represent the
same features of the white-chested bear, but in hypertrophied form
(still greater shortening in the facial parts, and broadening of the
zygomatic arches posteriorly, posterior broadening of the braincase,
mainly at the expense of the occipital crest, the further reduction
of tubercles, crests etc. of the skull, etc.). The systematic signifi-
cance of these peculiarities may, to the fullest degree, be understood

only with revision of all species (У.Н.).

Geographic Distribution

Forested regions of middle and southern parts of East Asia, the
Himalayas, Afghanistan and Baluchistan.

[ВАНА

Fig. 174. Reconstructed northern limit of occurrence of white-chested
(black) bear Ursus (Selenarctos) thibetanus G. Cuv. in USSR (V.G.
Heptner).

484

121

Geographic Range т the Soviet Union

The range in the USSR comprises the extreme northeastern part of
the species range and occupies a very small part of the country—
Ussuri territory and the region adjacent to the Amur river on the
north.

The northern border of the range runs from Innokenti Bay on
the coast of the Sea of Japan (about 50 km south of Sovietsk
Gavan) southwest to the region of chief heights (crest) of the
Sikhote-Alin’, crossing it at the sources of the Samarga. From this
place, the boundary directs itself to the north, through the middle
course of the Khor, Anyui and Khungari rivers and comes to the
shore of the Amur, crossing it at the level of the Gorin r[iver]
mouth. Along this river [Amur], the bear has been noted as far as
51° N. Lat. (Bagetor camp near Lake Evoron; K.A. Vorob’ev).

Thence, the boundary runs at some distance from the Amur
valley along its left bank, being directed towards the southwest, and
passes to the north of Lake Bolon’ and the point of juncture of the
Kur and Tunguska. The white-chested bear is encountered in the
lower course of Urmi. Farther to the west, it occurs north of the
bend of the Amur, between the mouth of Ussuri and 130°30’ Е.
Long. and the Tyrma river on the north, and particularly on the
southern slopes of Dur and Shuki-Pokta ranges. Thence, the bound-
ary of the range, crossing the southern spurs of the Bureinsk
mountains—the Vand and Aagar-Aul ranges—exits into northeastern
China along the southern slopes of the Malyi Khingan [range].
Probably, in some places, the bear penetrates somewhat farther
north than the above-mentioned line. It is absent on Sakhalin.”

In Ussuri territory, the black bear is not found everywhere,
being restricted to the broad-leaved forests which are connected
with the forests of the so-called Manchurian type; it avoids the
open areas and the high parts of Sikhote-Alin’

“The assertion of Sowerby (1923), concerning the occurrence of this
species in Kamchatka, used by several authors, is without any foundation.
Range according to Shrenk, 1858; Maak, 1861; Radde, 1862; Emel’yanov, 1927;
Ognev, 1931; the data of K.A. Vorob’ev and, primarily, according to Bromlei,
1956.

Fig. 175. Species range of the white-chested bear, Ursus (Selenarctos)
thibetanus G. Cuv. (reconstructed) (V.G. Heptner).

Geographic Range outside the Soviet Union

The (reconstructed) range outside the USSR occupies Baluchistan
(Suleiman mountains, but also Makran coast), the Hindu Kush [moun-
tain] system (southern) in Afghanistan (D. Povol’nyi), Punjab,
Kashmir, Himalayas within the boundaries of Nepal, Sikkim and
Bhutan, and farther east—the southern slope, but locally, and appar-
ently, also the northern side within the boundaries of Tibet. The

485

723

range includes Assam, Burma, Indochina Peninsula, Malay* Penin-
sula south to Mergui (12°30’ М. Lat.) or to the northern boundary
of the southern part of the Malay* Peninsula (about 7—7°30° М.
Lat.), southeastern and eastern China, including Sichuan and Gansu,
the eastern part of northeastern China, Hainan, Taiwan and the
Japanese Islands (absent on Ryukyus).”°

At the present time, the range has changed considerably, ex-
cept in the south and north, in the eastern half of China where the
bear is absent; it exterminated in a series of places, and also in
Japan (V.H.).

Geographic Variation

The described species has been given about 15 names. In the past,
7—8 races usually have been accepted, but this is, apparently, too
many, especially if the small range of the species is considered.
Some races were evidently described without sufficient basis.
However, several are, apparently, well characterized. It is neces-
sary to revise geographic variation in the species.

There is one subspecies within the boundaries of our country:

Ussuri white-chested bear, U. (S.) Е ussuricus Heude, 1901.

For description, see above.

Found in Ussuri and Amur territories.

Outside USSR—in northeastern China (former Manchuria),
Korean Peninsula, and northern parts of eastern China.

The Ussuri bear is a well-defined form, distinguished from
southern [forms], not to mention the insular ones, by its pure black,
dense and long fur, and larger dimensions. For Kashmir bears, it is
true, lengths of up to 202 cm and weight up to 181 kg have been
recorded; however, the accuracy of these measurements is doubt-
ful. In any event, animals with greatest length of skull more than
306 mm are not known from India, Pakistan, and Burma (Pocock,
1941). Therefore, the largest greatest length of skull of an Indian
specimen is less than the average for this measurement in Ussuri
bears. The largest recorded condylobasal length of skull of animals

*Russian original erroneously reads “Malakka’”—Sci. Ed.

S°Older authors refer to its occurrence the entire archipelago (Temminck,
1847), and more recent (Ellerman and Morrison-Scott, 1951), only in Honshu
(Hondo), Kyushu and (?) Shikoku.

486

724

from Hubei and Sichuan (275.0 mm; Allen, 1938), are also less than
the average of our bears.

Outside the borders of our country, the following forms are
usually accepted: 1) U. (S.) ¢t. gedrosianus Blanf. 1877—
Baluchistan; 2) U. (S.) t. laniger Pocock, 1932—Kashmir, Punjab,
Afghanistan; 3) U. (S.) Е. thibetanus Cuvier, 1823—from Nepal
through Assam to Burma, Siam and Vietnam inclusive; 4) Ц. (5.)
t. melli Matschie, 1922—from Yunnan to Fujian, Hainan; 5) U. (S.)
t. mupinensis Heude, 1901—Sichuan, Shanxi, Hubei; 6) Ц. (S.) ¢.
japonicus Schlegel, 1857—islands of Honshu (Hondo), Kyushu (?),
Shikoku; 7) U. (S.) Е. formosanus Swinhoe, 1864—Taiwan.

The small form, japonicus is very well-marked; the skull of a
2.5 year old female from Japan (Zoological Museum, Moscow
University) has a greatest overall length of 226 mm, condylobasal
length—198 mm, and zygomatic width—141 mm. As mentioned,
ussuricus differs, apparently, from the well-validated insular form,
formosanus, and the extreme southwestern form, gedrosianus.
Distinctiveness of /aniger from thibetanus and the validity of
mupinensis are highly doubtful (V.H.).

Biology

Population. There are no accurate data on numbers. It is more
numerous in southern Primor’e, and is rarely met with in the north-
ern spurs of Sikhote-Alin’ range. Within the limits of the Tachin-Chtan
range, for each 20 white-chested bears, one brown bear is found.
In Sikhote-Alin’ preserve, for one white-chested bear, 20 brown
bears are found (Bromlei, 1956; Yu.A. Salmin and V.D. Shamykin).

Habitat. The white-chested bear prefers oak-broad-leaved and
nut-pine-broad-leaved forests of the Manchurian type, and rarely
occupies fir-spruce taiga and high-montane birch krummholz and
montane balds. They avoid forests of the Okhotsk type.

They are more often encountered in river valleys and adjacent
slopes.

They set off to the mountain tops only when there are crops of
“nutlets” of nut-pines and red billberries.

Food. In April and May, before the appearance of green veg-
etation, the white-chested bear feeds at the expense of nut-pine
“nutlets” and acorns of the previous year. In case of crop failure,

725

Fig. 176. Broad-leaved and coniferous broad-leaved forests of “Kedrovaya

Pad’ ” preserve (southern Primor’e). Typical place of habitation of white-

chested bear; here also are spotted deer, roedeer, wild bear, yellow-throated
marten, leopard. June, 1963. Photograph by A.G. Pankrat’ev.

it searches in river valleys for nuts of Manchurian filbert [hazel-
nut], and captures insect larvae in rotten logs. Nursing females do
not go far from dens, feeding also on young sedge and birch sap;
the latter, possibly, favors removal of the “plug” from the rectum.

From mid-May and in all of June, food is monotonous—stems of
coltsfoot, cow parsnip, angelica, sedge, ovaries of oak, leaves of
Amur cork tree and Manchurian walnut; from the end of June, they
add the fruits of honeysuckle, searching in old burns along sunny
slopes.

All of July, August and part of September, its main food con-
sists of fruits of bird-cherry which the bears eat while climbing
trees. While eating, they bend and break branches beneath them-
selves, as a result of which places are formed in crotches of
trunks—“nests”; such “nests” also originate in oaks with feeding

487

488

726

999

preserve—biotope
of white-chested bear. Southern Primor’e. 20 September 1964. Photograph
by A.G. Pankrat’ev.

Fig. 177. Black-fir-hornbeam forest in “Kedrovaya Pad

by bears on acorns. In August, food also includes Amur jack-in-
the-pulpit and its bulbs, iris, ovaries of oak and other plants.

In September, to bird-cherry they add acorns (often still un-
ripe), nut-pine cones, actinidia vine, Amur grape and rarely, still-green
filbert nuts; green plants disappear from the diet. At the end of
September, nut-pine “nutlets” and acorns become the principal
food and remain so in harvest years until retirement to the den.

In years of crop failure, white-chested bears descend to river
valleys and springs and, before retiring into dens feed on Manchu-
rian walnuts, Amur grapes, actinidia and rarely on berries of Amur
cork tree. Under these conditions, they almost never climb trees.
Predatoriness is not typical for the white-chested bear, even with
no crops of their principal foods. In rare cases, bears feed on
carrion and fish which die after spawning (sima salmon) [O. masu]
but, as a rule, they do not eat fish.

727

487

Fig. 178. Burrow of a hibernating Siberian chipmunk destroyed by white-
chested bear with goal of taking store of nut-pine “nutlets”. “Кедгоуауа
Pad’ ” preserve. 5 November 1964. Photograph by A.G. Pankrat’ev.

Home range. There is no information. In spring, the nursing
females do not move farther than 1—1.5 km from the den.

Burrows and shelters. In winter, the white-chested bears re-
tire to hollows in old trees, usually Maximovich poplar, and Chosenia,
linden and oak. They also frequently hibernate in caves and rocks.
If disturbed, or driven from dens, they occasionally lie down in
hollows without shelter, on the ground at the base of a tree, and
beneath roots.

Of 42 dens, 55% were situated in hollow [trees] (linden—22%,
poplar—26%, oak—7%), in rocks—38%, and directly on the ground—
7% (Bromlei, 1956). The percentage of dens in rocks is, apparently,
inflated, because they are easily discovered. Bears occupy dens for
many years in a row. Animals grossly fattened—bears well fed for
winter—may not be able to pass through entrances of hollows, and
therefore, large bears more often retire in rocks in [good] food
years.

728

488 Fig. 179. Hollow in Amur linden, in which there were wild Indian bees.
torn apart by white-chested bear to obtain honey. “Kedrovaya Pad’ ”
preserve, Southern Primor’e. November 1962. Photograph by A.G.

Pankrat’ev.

Daily activity and behavior. The daily cycle of activity is not
known. Behavior of the white-chested bear differs from the brown
in that it often climbs trees and feeds on them. They also climb in
thickets of vines.

Hibernation and winter sleep. In autumn, the white-chested
bears accumulate fat more quickly and simultaneously than brown,
and retire to dens somewhat earlier (V.D. Shamykin, Yu.A. Salmin).
With the onset of cold, these bears are rarely met with, and at the
end of October—-beginning of November, even before snowfall, they
have disappeared. Cases of delay in retirement and arousal

489

729

(“shatun’’) have not been noted in this species (Sikhote-Alin’ range).
However, in Tachin-Chtan range, white-chested bears retire later
than brown (Bromlei, 1956). Individual animals were seen in the
marine littoral zone at the end of December. At the beginning of
winter, they sleep lightly, later on—heavily. Females with cubs do
not leave the hollows even when smoked. Exit from the den in
spring occurs later than in the brown—in the second half of April,
when the snow has already melted. Therefore, white-chested bears
do not experience sharp spring hunger. Near the sea and on the
southeastern slopes of the Sikhote-Alin’, bears wake up earlier than
in the interior of the mainland; those lying in hollows exit later than
those lying in rocks; females with cubs get up last of all up. Some-
times, after a heavy late snowfail in April, they again retire. Animals
driven from dens sometimes go to sleep again—even in the open at
the base of a tree trunk; cases are known when a female bear in
such a position carried the cubs and hid them in her abdominal wool
(Bromlei, 1956). In the den, they thin little during the period of
winter sleep, and only the nursing females lose fat; fat is primarily
consumed in the period of spring food shortage.

Seasonal migrations and transgressions. In searching for
food, bears undertake movements of about 200—350 km. They oc-
cur more frequently than in the brown bear, but for lesser distances,
and they are closely connected with yield and distribution of the
main plant foods (nuts, acorns).

Reproduction. In Sikhote-Alin’, rut in the white-chested bear
is initiated earlier than in the brown—from the middle of June to the
middle of August. Accordingly, birth takes place earlier: nursing
females were taken from the middle of January (Yu.A. Salmin, and
V.D. Shamykin). Mating was noticed in the region of the Tachin-
Chtan range from the first days of June to the end of July (wide
paths, flattened areas of trampled grasses with 10—15 m diameter).
The first signs of pregnancy (swelling of uterine horns to 15—22
mm) are observed in October. By the end of December, the weight
of the embryo reaches 75 g. All dates of reproduction have an
amplitude [of variation] of 2 months (Bromlei, 1956). Barrenness is
a frequent phenomenon (for 1 pregnant female—2 barren).

The female gives birth to 2 offspring, rarely 3, beginning from
the end of December (very rarely) to the middle of February,
occasionally to the beginning of March, but most often from the
middle of January to the middle of February.

730

Fig. 180. Hollow of wild Indian bees in Daurian birch, torn apart by white-
chested bear. “Kedrovaya Pad’ ” preserve, southern Primor’e. March 1965.
Photograph by A.G. Pankrat’ev.

Growth, development, and molt. The bear-cubs are born blind,
weighing 300—450 g, and differ from the brown in the greater length
490 of their feet and claws. They grow slowly, reaching in May only
2.5 kg (rarely 4.0 kg). Sometimes, “runts” occur, weighing at the
end of summer 6—7 kg (diseased or born late). The first month after
her departure from the den, the she-bear boldly defends the cubs
(April—first half of May). In June, after driving the cubs into trees,

she flees (Bromlei, 1956).
Enemies, diseases, parasites, mortality, competitors, and
population dynamics. Tigers and wolves may be enemies of the

731

Fig. 181. Clearing with tall grasses in the forest of “Kedrovaya Pad’ ”
preserve, southern Primor’e. Place of feeding of white-chested she-bear
with cubs. August 1963. Photograph by A.G. Pankrat’ev.

white-chested bear, and for cubs, also lynx. The tiger can cope
with a bear weighing 60 kg.

The white-chested bear rarely comes across the brown, be-
cause they primarily restrict themselves to different altitudes and
occupy different stand-types. They are found frequently together
only in certain stand-types (foothills, seashore) in years of acorn
harvest.

In April (mainly), they are attacked by ticks; in August, lice are
observed. Infestation with intestinal worms is negligible (Bromlei,
1956). |

The frequency of occurrence of the white-chested bear in the
region of the Tachin-Chtan range is not constant; changes are
directly related to the autumn food crop (nuts, acorns). In 1944 in
this region, following a doubling of the yield of acorns and nut-pine
“nutlets”, a considerable concentration of bears emerged.

491

792

Field characteristics. Signs of activity of the white-chested
bear differ from those of the brown in the presence of “arbors”
of twigs in the forks of trees, and in claw marks on the bark [of
trees] to a height of more than 2.5 m.* Animals frequently climb
in thickets of vines, and dig a lot in the ground (P.Yu.).

Fig. 182. “Nest” made of broken branches left in the tree top after feeding by

white-chested bear on the fruits of a large tara vine (actinidia). “Kedrovaya

Pad’” preserve, southern Primor’e. September, 1958. Photograph by
A.G. Pankrat’ev.

*Lapsus ? Brown bears are more likely to leave claw marks at a height

of “more than 2.5 m. than are the much smaller white-chested bear—Sci.
Ed.

183

Practical Significance

The white-chested bear yields skin, meat and fat of greater value
than does the brown bear. It does not belong among the predators
injurious to game or domestic animals. Dens of the white-chested
bear are easily discovered, and therefore the animal is easily exter-
491 minated; it is in need of protection. Its utilization must be strictly

Fig. 183. Marks of an Asian black bear on a white-barked fir. Marks made
in spring. Western shore of Amur Gulf, southern Primor’e. June, 1962.
Photograph by A.G. Pankrat’ev.

regulated. In the case of den-hunting, tree hollows are often de-
stroyed, which reduces the number of shelters necessary for
overwintering.

They are hunted with dogs (P.Yu.).

ie
ae

yy
x
И
№
И

en
Lae

И
ИЯ

ыы )
a

Other Books Inter

MAMMALS GF THE SOVIET UNION
Volume II, Part 2

This volume, is part of a three-volume monograph, and is a continuation,
of Volume II, Part 1, which was devoted to sea cows and carnivores. It
contains species descriptions of terrestrial carnivores and detailed informa-
tion on their external morphology, skull, body measurements and other
data, affinities with other species, geographic distribution in the historic
past and today, geographic variation, practical significance, and biology.
Descriptions are presented for orders, families, and genera, and keys ae
for their identification.

The book is richly illustrated with photographs, sketches, and colored
illustrations by the famous wild life painter, А.М. Komarov, and the
-zoologist-artist, ММ. Kondakov.

The results of original scientific studies are published here for the first
time, providing readers a vast wealth of material heretofore unknown. This
_ work is intended for teachers and students of faculties of biology and
geography in universities, as well as pedagogic, agricultural, and forest |
institutes, similar organizations, workers engaged in game, fur, forest and

_ fish trades; those interested in the conservation of nature, and all persons

interested in — and nature study.

MAMMALS OF THE SOVIET UNION |
ee Ц, Part 3

This volume is fourth book of Marion of the Soviet Union, representing
the third part of the second volume; it is devoted to descriptions of the
orders of Soviet aquatic mammals — pinnipeds (Pinnipedia) and, in part,
cetaceans (Cetacea), toothed whales (Odontoceti). In the sequence of
descriptions from the “higher” to the “lower” orders adopted in this series,
‘pinnipeds should have preceded carnivores, i.e., should have appeared in
the second book. The grouping of the orders ata higher level is given after
G.G. Simpson (1945). The sequence of genera and species within the
orders has been retained as before, i.e., from the less specialized to the
_ more specialized.
_ Т№е order of pinnipeds or seals (Pinnipedia) is described in this
volume. The cohort of ungulates and carnivores represented in Soviet fauna
by orders of artiodactyls, and perissodactyls, sirenians, carnivores, and
pinnipeds thus came to an end and the cohort of whales (Mutica) com-
-mences. The toothed whales are described in this volume.

While it has not always been possible to maintain a totally uniform
description of the genera and species as in the volumes already published,
in spite of every effort to do so, the sequence has been adhered to, with

_ зоте exceptions, in the case of Pinnipedia. But, it was impossible to use

same format in describing the toothed whales (Odontoceti). Only a brief

morphological description has been given for many species, which is more
or less adequate for identifying the species. General information on their
distribution and fragmentary biological data are also given. In some cases
_ the total absence of such information is indicated. =

Internet Archive Audio

Featured

Top

Images

Featured

Top

Software

Featured

Top

Texts

Featured

Top

Video

Featured

Top

Mobile Apps

Browser Extensions

Archive-It Subscription

Save Page Now

Full text of "Mammals of the Soviet Union"

See other formats